ENCYCLOPEDIA OF

FOOD GRAINS
Second Edition
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 ENCYCLOPEDIA OF
FOOD GRAINS
Second Edition
VOLUME 1
THE WORLD OF FOOD GRAINS
EDITORS
COLIN WRIGLEY
University of Queensland, Brisbane, Australia

HAROLD CORKE
University of Hong Kong, Hong Kong, Peoples Republic of China

KOUSHIK SEETHARAMAN{

JON FAUBION
Kansas State University, Manhattan KS, USA

{
deceased

AMSTERDAM BOSTON HEIDELBERG LONDON

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Academic Press is an imprint of Elsevier
Academic Press is an imprint of Elsevier
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First edition 2004 (as Encyclopedia of Grain Science)

Copyright 2016 Elsevier Ltd. All rights reserved

The following articles are US government works in the public domain and are not subject to copyright:

Grain Crops, Overview; Maize Overview.

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SET ISBN 978-0-12-394437-5

VOLUME 1 ISBN 978-0-12-803537-5
VOLUME 2 ISBN 978-0-12-803536-8
VOLUME 3 ISBN 978-0-12-803535-1
VOLUME 4 ISBN 978-0-12-803538-2

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CONTENTS

Preface xvii
Tributes to Former Co-Editors xix
Editors Biography xxiii
Editorial Advisory Board xxv
Contributors xxvii

VOLUME 1

THE WORLD OF FOOD GRAINS 1

The Basics 1
The Grains that Feed the World 1
CW Wrigley, H Corke, and J Faubion

Course Structures: Based on EFG Articles 13

H Corke

The Grain Crops: An Overview 16

RA Graybosch

The Grain Chain: The Route from Genes to Grain-Based Products 22

CW Wrigley

Taxonomic Classification of Grain Species 31

LA Morrison and CW Wrigley

Grain: Morphology of Internal Structure 41

H Corke

Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 51
J Wyatt

The Cereal Grains 73

An Overview of the Family of Cereal Grains Prominent in World Agriculture 73
CW Wrigley

Cereals: Domestication of the Cereal Grains 86

LA Morrison

Maize: Overview 99
MP Scott and M Emery

Wheat: An Overview of the Grain That Provides Our Daily Bread 105
CW Wrigley

v
vi Contents

Durum Wheat: Overview 117

GP Kadkol and M Sissons

Rice: Overview 125

BO Juliano

Wildrice, Zizania: Overview 130

EA Oelke and RA Porter

African Rice (Oryza glaberrima): A Brief History and Its Growing Importance in Current
Rice Breeding Efforts 140
JT Manful and S Graham-Acquaah

Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses 147
AM Stanca, A Gianinetti, F Rizza, and V Terzi

Sorghum Grain, Its Production and Uses: Overview 153

A Cruickshank

Rye Grain: Its Genetics, Production, and Utilization 159

HD Sapirstein and W Bushuk

Triticale: Overview 168

BJ Furman

Oats: Overview 173

PK Zwer

Coix: Overview 184

H Corke, Y Huang, and JS Li

Millet Pearl: Overview 190

JRN Taylor

Millet Minor: Overview 199

GK Chandi and GA Annor

Teff: Overview 209

G Bultosa

The Oilseeds 221

Oilseeds: Overview 221
RJ Mailer

Soybean: Overview 228

K Liu

Canola: Overview 237

VJ Barthet

Cottonseed: Overview 242

E Hernandez

Sunflower: Overview 247

GJ Seiler and TJ Gulya

Overview of the Oilseed Safflower (Carthamus tinctorius L.) 254

C Hall III

Linseed: Overview 259

S Cloutier

The Legumes and Pseudocereals 265

Grain Legumes and Their Dietary Impact: Overview 265
TE Michaels
 Contents vii

Pseudocereals: Overview 274

RJ Fletcher

Lupin: Overview 280

DS Petterson

Amaranth: Overview 287

H Corke, YZ Cai, and HX Wu

Beans: Overview 297

SK Sathe

Buckwheat: Overview 307

YZ Cai, H Corke, D Wang, and WD Li

Chickpea: Overview 316

EJ Knights and KB Hobson

Pea: Overview 324

TN Khan, A Meldrum, and JS Croser

Peanuts: Overview 334

RCN Rachaputi and G Wright

Quinoa: Overview 341

SA Valencia-Chamorro

Grains Around the World 349

Grain Production and Consumption: Overview 349
T Beta and C Isaak

Grain Production and Consumption: Africa 359

JRN Taylor

Production and Consumption of Grains: India 367

BS Khatkar, N Chaudhary, and P Dangi

Grain Production and Consumption: China and South-East Asia 374

H Corke and YZ Cai

Grain Production and Consumption: Europe 383

MG Lindhauer

Grain Production and Consumption: Cereal Grains in North America 391

JA Fox and LL Ward

Grain Production and Consumption: Oilseeds in North America 401

PBE McVetty, OM Lukow, LM Hall, I Rajcan, and H Rahman

Grain Production and Consumption in Oceania: Australia and Pacific Countries 409
CW Wrigley and RJ French

Grain Production and Consumption: South America 420

OR Larroque and JC Catullo

Scientific Organizations Related to Grains 429

Research Organizations of the World: CGIAR 429
JH Skerritt

Research Organizations of the World: Europe and North America 436

JH Skerritt

Research Organizations of the World: Asia-Pacific, CentralSouth America, and AfricaMiddle East 444
JH Skerritt
viii Contents

Research Organizations of the World: Global Trends and the Commercial Sector 451
JH Skerritt

Scientific Societies Associated with Grain Science 457

SC Nelson and CW Wrigley

VOLUME 2

NUTRITION AND FOOD GRAINS 1

Food Grains and Well-Being 1

Functional Foods: Overview 1
G Bultosa

Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics 11

G Bultosa

Nutrition: Soy-Based Foods 17

AM Fehily

Food Grains and the Consumer 23

Grains and Health: Misinformation and Misconceptions 23
DB Sheats and JM Jones

Consumer Trends in Grain Consumption 29

JM Jones and DB Sheats

Cultural Differences in Processing and Consumption 35

AMR Hayes and JM Jones

Fortification of Grain-Based Foods 43

CM Rosell

Genetically Modified Grains and the Consumer 50

A Mathiowetz and JM Jones

Labelling of Grain-Based Foods 57

JM Jones

Grains and Health 63

R Korczak, D Hauge, B Maschoff, L Marquart, P Jacques, R Lindberg, and R Menon

Food Grains: Intolerance, Allergy and Diseases 72

Cereal Allergens 72
AS Tatham

Nutrition: Beriberi, A Deficiency Related to Grains 77

KJ Carpenter

Celiac Disease 83
P Koehler, H Wieser, and KA Scherf

The Gluten-Free Diet 91

V Zevallos and I Herencia

Proteins 98
The Protein Chemistry of Cereal Grains 98
F Bekes and CW Wrigley

The Protein Chemistry of Dicotyledonous Grains 109

PR Shewry
 Contents ix

Protein Synthesis and Deposition 115

PR Shewry and P Tosi

Nitrogen Metabolism 123

CA Atkins

Nitrogen in Grain Production Systems 129

XM Zhou, BL Ma, and DL Smith

The Enzymes Associated with Food Grains and Their Functions in Processing 138
DJ Mares, K Mrva, and GB Fincher

Proteomic Analysis Relevant to Cereal Grains 146

A Juhasz, R Haraszi, F Bekes, DJ Skylas, and CW Wrigley

Thiolomics of the Gluten Protein Network of Wheat Dough 154

F Bonomi and S Iametti

Carbohydrates 161
Carbohydrate Metabolism 161
RN Chibbar, S Jaiswal, M Gangola, and M Baga

Starch: Chemistry 174

FL Stoddard

Starch: Synthesis 181

A Regina, S Rahman, Z Li, and MK Morell

Starch: Starch Architecture and Structure 190

V Vamadevan and Q Liu

Starch: Analysis of Quality 198

A Gunaratne and H Corke

Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 208

GB Fincher

Grains other than Cereals: Non-starch Polysaccharides 224

L Ramsden

Resistant Starch and Health 230

A Evans

Health Effects of b-Glucans Found in Cereals 236

SM Tosh and S Shea Miller

Glycemic Index, Glycemic Load, and Their Health Benefits 241

D Dan Ramdath

Fats 248
Lipid Chemistry 248
L Day

Healthy Fats and Oils 257

SM Ghazani and AG Marangoni

Bioactives and Toxins 268

Bioactive Compounds in Wheat Bran 268
HD Sapirstein

Bioactives: Antioxidants 277

T Beta and KG Duodu
x Contents

The Antinutritional Components of Grains 283

CK Madsen and H Brinch-Pedersen

Mycotoxins 290
MT Fletcher and BJ Blaney

Grain Composition and Analysis 297

The Composition of Food Grains and Grain-Based Products 297
CW Wrigley

Standardized Test Methods for Grains and Grain-Based Products 300

AR Bridges and CW Wrigley

Units of Grain Science and Trade: Equivalence between the US, Chinese, and Metric Units 308
W Huang and CW Wrigley

VOLUME 3

GRAIN-BASED PRODUCTS AND THEIR PROCESSING 1

Wheat-Based Foods 1
Cereals: Overview of Uses: Accent on Wheat Grain 1
CF Morris

Breads 8
TR Moore

Flatbreads of the World 19

KJ Quail

Sour Dough Technology 25

SP Cauvain

Cakes, Muffins and Bagels 30

DE Ortiz

Cookies, Biscuits and Crackers: Formulation, Processing and Characteristics 37

SP Cauvain

Cookies: A Diverse Family of Baked Goods 44

S Zydenbos, V Humphrey-Taylor, and CW Wrigley

Wafers: Methods of Manufacture 51

K Tiefenbacher and CW Wrigley

Cookies and Crackers: Commercial Production 59

LC Paulson and CW Wrigley

Noodles: Starch 64
LS Collado and H Corke

Noodles: Asian Wheat Flour Noodles 72

GB Crosbie and AS Ross

Pasta 79
M Sissons

Tortillas 90
LW Rooney and SO Serna-Saldivar

Non-wheat Foods 97
Maize: Foods from Maize 97
SO Serna-Saldivar
 Contents xi

Rice: Chinese Food Uses 110

S Lu

Sorghum: Utilization 116

RD Waniska, LW Rooney, and CM McDonough

Soybean: Soy-Based Fermented Foods 124

DK OToole

Soybean: Soymilk, Tofu, and Okara 134

DK OToole

Food-quality Testing 144

The Application of Sensory Science to the Evaluation of Grain-Based Foods 144
LM Duizer and SB Walker

Noodles: Testing for Quality 154

VA Solah and GB Crosbie

Pasta: Quality Testing Methods 161

A Marti, MG DEgidio, and MA Pagani

Rice: Eating Quality 166

JS Bao

Beverages from Grains 176

Fermentation: Origins and Applications 176
BJB Wood

Fermentation: Foods and Nonalcoholic Beverages 183

JRN Taylor

Beverages: Distilled 193

GH Palmer

Beverages: Asian Alcoholic Beverages 206

DK OToole

Non-food Products from Grains 215

Cereal Grains as Animal Feed 215
JL Black

Pet Foods 223

RCE Guy

Utilization of the Whole Cereal Plant to Maximize its Value 228

L Munck

Fuel Alcohol Production 235

YN Guragain, KV Probst, and PV Vadlani

Biodiesel 245
Y Yan

Processing of Grains 251

Evaluation of Wheat-Grain Quality Attributes 251
CF Morris

Grain Quality Attributes for Cereals Other than Wheat 257

CF Morris
xii Contents

Cereals: Breakfast Cereals 262

EF Caldwell, JD McKeehen, and RS Kadan

Extrusion Technologies 268

RCE Guy

Starch: Uses of Native Starch 274

JW Lawton

Starch: Modification 282

JN BeMiller

b-Glucans: Measurement and Processing 287

MS Izydorczyk

Wheat Processing 299

Milling and Baking: History 299
CE Walker and WD Eustace

Wheat: Dry Milling 307

AK Sarkar and JE Dexter

Wet Milling of Wheat 320

R Velicogna and S Shea Miller

Oven Technologies 325

CE Walker

Bakeries: The Source of Our Unique Wheat-Based Food, Bread 335

JE Bock, CW Wrigley, and CE Walker

Analysis of Dough Rheology in Breadmaking 343

B Dobraszczyk

Frozen Dough 354

D Domingues and C Dowd

Refrigerated Dough 359

D Domingues, C Dowd, and W Atwell

Chemistry of Cake Manufacturing 367

TS Palav

The Gluten Proteins of the Wheat Grain in Relation to Flour Quality 375
F Bekes, MC Gianibelli, and CW Wrigley

Ultrastructure of the Wheat Grain, Flour, and Dough 384

S Grundas and CW Wrigley

Cereal Food Production with Low Salt 396

L Day

Baked Product Staling: Mechanisms, Determinations, and Anti-staling Strategies 403

A Goldstein and K Seetharaman

Gluten and Modified Gluten 408

IL Batey and W Huang

Snack Foods: Processing 414

MN Riaz

Barley, Rice and Maize Processing 423

Barley: Malting 423
L MacLeod and E Evans

Barley: Milling and Processing 434

MS Izydorczyk and JE Dexter
 Contents xiii

Rice Processing: Beyond the Farm Gate 446

L Pallas

Oil from Rice and Maize 453

JS Godber

Maize: Dry Milling 458

KD Rausch and SR Eckhoff

Maize: Wet Milling 467

KD Rausch and SR Eckhoff

Oilseed and Legume Processing 482

Soybean: Soy Concentrates and Isolates 482
C-Y Ma

Soybean: Processing 489

T Wang

Canola: Processing 497

JK Daun and EH Unger

VOLUME 4

THE PRODUCTION AND GENETICS OF FOOD GRAINS 1

Grain Marketing and Grading 1

Wheat: Grading and Segregation 1
RL Cracknell and RM Williams

Wheat: Marketing 9
RL Cracknell and RM Williams

Barley: Grading and Marketing 16

A MacLeod, M Edney, and MS Izydorczyk

Soybean: Grading and Marketing 25

EG Hammond, LA Johnson, and PA Murphy

Identification of Varieties of Food Grains 29

DM Miskelly and CW Wrigley

Grain Harvest, Storage and Transport 42

Wheat: Harvesting, Transport, and Storage 42
S Grundas and CW Wrigley

Barley: Harvesting, Storage, and Transport 50

RJ Henry

Sorghum: Harvest, Storage, and Transport 54

T Beta, M Chisi, and ES Monyo

Canola: Harvest, Transport, and Storage 62

JJ Mayko

Cereals: Grain Defects 68

SA Barton

The Nature, Causes, and Control of Grain Diseases in the Major Cereal Species 74
RA McIntosh, PM Williamson, and CW Wrigley

Contaminants of Grain 83
J McLean and CW Wrigley
xiv Contents

Chemicals for Grain Production and Protection 99

PC Annis

Handling from Farm to Storage Terminal 105

D Richard-Molard and CW Wrigley

Stored Grain: Invertebrate Pests 110

PC Annis

Postharvest Operations for Quality Preservation of Stored Grain 117

F Fleurat-Lessard

Stored-Grain Pest Management 126

F Fleurat-Lessard

Agronomy of Grain Growing 140

Implication of Climate Changes 140
S Ceccarelli

Sustainable Grain Production and Utilization 144

L Munck

Organic Grain Production and Food Processing 154

P Gelinas and C David

Precision Agriculture 162

EC Leonard

Plants: Diseases and Pests 168

SA Barton

Wheat: Agronomy 176

GM Paulsen, JP Shroyer, and KJ Shroyer

Barley: Agronomy 186

RD Horsley and M Hochhalter

Maize: Agronomy 194

ED Nafziger

Sorghum: Production and Improvement Practices 201

WL Rooney

Canola: Agronomy 207

NJ Mendham and MJ Robertson

Chickpea: Agronomy 216

KHM Siddique and L Krishnamurthy

Lentil: Agronomy 223

T Nleya, A Vandenberg, FL Walley, and D Deneke

Lupin: Agronomy 231

RJ French

Field Pea: Agronomy 240

RJ French

Soybean: Agronomy 251

ED Nafziger

Natural Disease Control in Cereal Grains 257

WK Mousa and MN Raizada

Wheat: Biotrophic Pathogen Resistance 264

RF Park
 Contents xv

Necrotrophic Pathogens of Wheat 273

RP Oliver, K-C Tan, and CS Moffat

Breeding of Grains 279

Wheat Breeding: Exploiting and Fixing Genetic Variation by Selection and Evaluation 279
R DePauw and L OBrien

Barley: Genetics and Breeding 287

A Cuesta-Marcos, JG Kling, AR Belcher, T Filichkin, SP Fisk, R Graebner, L Helgerson, D Herb, B Meints, AS Ross,
PM Hayes, and SE Ulrich

Maize: Breeding 296

EA Lee and LW Kannenberg

Rice: Breeding 304

Qingyao Shu and Dianxing Wu

Canola/Rapeseed: Genetics and Breeding 311

PBE McVetty and RW Duncan

Lentil: Breeding 317

W Erskine, A Sarker, and S Kumar

Lupin: Breeding 325

LC Trugo, E von Baer, and D von Baer

Soybean: Germplasm, Breeding, and Genetics 333

RG Palmer and T Hymowitz

Genetics of Grains 343

Development of Genetically Modified Grains 343
L Privalle

Detection of Genetically Modified Grains 350

RD Shillito

Genomics of Food Grains 360

RJ Henry

Genome Mapping 365

VK Tiwari, JD Faris, B Friebe, and BS Gill

Wheat Genetics 376

RA McIntosh

Wheat Genetics and Genomics 382

E Akhunov

Biotic Stress Resistance Genes in Wheat 388

J Singla and SG Krattinger

Abiotic Stress Genes and Mechanisms in Wheat 393

SJ Roy, NC Collins, and R Munns

Rice: Genetics 398

JS Bao and H Corke

Maize: Genetics 407

EA Lee

Maize: Quality Protein Maize 420

JS Li and SK Vasal

Maize: Other Maize Mutants 425

MH Blanco, H Yangcheng, and J Jane
xvi Contents

Appendix 1. Nutrient-Composition Tables for Grains and for Grain-Based Products 433
SF Schakel, N Van Heel, and J Harnack

Appendix 2. Lists of Standardized Testing Methods for the Analysis of Grain and Grain-Based Foods 450
AR Bridges and CW Wrigley

Appendix 3. Grains, Foods, and Ingredients Suiting Gluten-Free Diets for Celiac Disease 460
JM Jones, V Zevallos, and CW Wrigley

Glossary 467
Index 491
PREFACE

The grain-producing plants are essential to life on earth. They are the primary means by which sunlight, the
primary source of the earths energy, is captured via photosynthesis to turn carbon dioxide and water into
carbohydrates and thus into the wider range of the biochemicals needed by all life forms.
The summary is simple:
The suns energy CO2 H2O
#
Grain-bearing plants
which provide:
Food, Fats, Fuel and Fiber for ourselves
Feed, Forage, Fodder for our animals
Food for Fish Farms

Directly, grains supply over half of humanitys energy and protein.

Indirectly, grains contribute to our food supply via the feeding of grains and grain by-products to animals,
birds and fish.
Grains are increasingly a renewable source of fuels (ethanol and oils).
Grain species, such as cottonseed, contribute fiber for our clothing.

Grains to Feed and Fuel our World

As its title indicates, the Encyclopedia of Food Grains concentrates on the food uses of grains, but details are also
provided about the wider roles of grains. Greater detail about the use of grains for animal feeding is already
available via the on-line Feedipedia encyclopedia (http://www.feedipedia.org/).
Worldwide, however, it is our food that is our obsession first and foremost. We see our health (or
otherwise) to be intimately related to our diet. Therefore nutrition is a major accent of this second edition,
occupying the whole of the second volume, following the first volumes description of the basic aspects of the
world of grains. The third and fourth volumes work their ways back up the value-added grain chain to describe
grain processing and production, thence to breeding and genetics.
The sequence of articles in the first edition (the Encyclopedia of Grain Science, 2004) followed an alphabetical
order. In contrast, the distinct ordering of articles in the second edition assembles fundamental concepts in
Volume 1 The World of Food Grains. Moving into the later volumes, greater complexity and erudition are
evident. Thus, unusual terms such as Transcriptomics, Aeciospore and Allelopathy are more likely to be
found in later volumes than in the first. In any case, reference to the glossary and the index should help in
elucidating such terms.

The First and Second Editions - Editors

This second edition is a considerable expansion on the content of the first edition, with its three print volumes
and 168 articles. Now, some twelve years later, the four print volumes of the Encyclopedia of Food Grains provide

xvii
xviii Preface

216 articles, many newly written for the new edition. Those articles that have been reprinted from the first
edition have been updated. In all cases, authors and reviewers are world renowned experts in their respective
fields.
This second edition has been more than three years in its development. In the early stages, the editors were
Harold Corke, Chuck Walker and Colin Wrigley (the three editors of the first edition). Sadly in 2012, Chuck
Walker passed away, leaving a gap that was difficult to fill.
Chuck was replaced by Koushik Seetharaman, who worked avidly in developing the concepts for this second
edition, including his contributions to a meeting of the editors and Elsevier staff in Sydney, January, 2013.
Sadly, Koushik suffered a heart attack in June, 2014. However, his contributions to the second edition during
this critical development stage were such that Elsevier decided that his name should remain as a co-editor.
Tributes to Professors Walker and Seetharaman are provided below.
In July, 2014, Professor Jon Faubion was appointed as a co-editor. Dr Faubion is the Charles Singleton
Professor of Baking and Cereal Science in the Department of Grain Science, Kansas State University. He is a
more than worthy replacement under such tragic circumstances.

Acknowledgement to All Involved

The production of an encyclopedia involves a wide range of experts with expertise in a diversity of areas of
knowledge. We acknowledge with grateful thanks the groups of contributors:
Members of the Editorial Advisory Board;
Authors who have shared their special knowledge and experience;
Elsevier staff who have provided extensive advice and administrative services, especially Donna de
Weerd-Wilson, Simon Holt and Rachel Gerlis (successive Supervisory Staff) and Gemma Tomalin,
Gemma Taft, Joanne Williams and Mark Harper (successive Content Project Managers).
The Editors
Harold Corke, Jon Faubion, Koushik Seetharaman, and Colin Wrigley
TRIBUTES TO FORMER CO-EDITORS

Professor Emeritus Charles (Chuck) E. Walker

Chuck Walker was one of the three editors of the first edition of Elseviers Grains
Encyclopedia (the Encyclopedia of Grain Science, 2004). He was again appointed
as an editor of this second edition. He participated in the planning stages for the
second edition until his untimely death on 26 April 2012. At the time of his
death, Chuck was a Professor Emeritus at the Department of Grain Science and
Industry at Kansas State University, Manhattan, KS, USA.
Charles Eugene Walker was born on 17 December 1936, in Winterset, Mad-
ison county, IA, USA, during the great depression. He was the eldest child of
Albert Cecil and Bessie Irene Headley Walker, poor farmers in Winterset, Mad-
ison county, IA, USA. He walked more than a mile to attend rural one-room
grammar schools in Warren county and graduated from high school in New
Virginia, IA, USA.
Better known as Chuck in later life, Dr Walker received a BS degree in
Chemical Engineering from Iowa State University, Ames, IA, USA, in May 1959. At a mixer party in the fall
of 1957, he met Shannon Rose Philp, a pretty freshman student from Loup city, NE, USA. This was just after a
summer spent working alone on a fire-lookout tower for the US Forest Service, in the wilds of Clearwater
National Forest, in Northern Idaho. They were married on 1 June 1958 and she became his wife of 54 years. She
later rewarded him with two sons, Alan Eugene and Tomas Charles, and they adopted a daughter, Bekianne.
Upon graduation, Chuck worked in flour milling research at the General Mills Central Research Laboratories
in the Minneapolis area from 1959 to 1962. He then accepted a fellowship to graduate school at North Dakota
State University in Fargo from 1962 to 1965, majoring in Cereal Chemistry. Chuck received his PhD in
Chemistry in the summer of 1966. For his PhD dissertation, he developed a micro brewery and studied barley
proteins and their influence on beer quality a rather ironic choice of topics, considering he remained a strict
teetotaler all his life.
While finishing his thesis, he joined the faculty of the North Dakota State College, Valley city, ND, USA
(19651974), teaching various physical science and chemistry courses to undergraduate students. Feeling the
pull of industry, he moved south to work for the Fairmont Foods Central Research Laboratory in Omaha, NE,
USA (19741980), where he did research, product development, and technical assistance on baking, pizza,
snack, and dairy products, eventually reaching the position of Associate Director of Research.
Dismemberment and sale of this Fortune-500 company provided him an opportunity to return to academia
at the University of Nebraska, Lincoln, NE, USA (19801987), where he developed a Cereal Technology
Program within the Department of Food Technology. He developed and taught various food science and
technology courses to both undergraduate and graduate students, and developed a vigorous research program.
He became Interim Department Head and Food Processing Center Director.
In December 1987, Chuck joined the Department of Grain Science and Industry at Kansas State University as
the Bakers National Educational Foundations Baking Science Professor. There he maintained a busy program

xix
xx Tributes to Former Co-Editors

in research, teaching, graduate student advising, publishing, traveling and consulting, specializing in baking
technology.
During his career, Chuck published more than 100 technical papers, and advised about 50 MS and PhD
graduate students. He was heavily involved in applied rheology (mixing and flow characteristics) of flour
products. In recognition of his commitment to international graduate students, Mr Roland Temme, TMCO,
Lincoln, NE, USA, established an international graduate student fellowship in his honor.
Chuck entered 50% phased-in retirement at age 65 in 2002 and retired in 2005, only to be hired back 1/
10-time afterward to assist with seminars and the departments centennial recognition in 2010. He continued
to support the departments activities on a volunteer basis.
Chucks research and consulting travels took him to more than a dozen countries, including many extended
trips to Australia, eventually earning permanent resident status in Australia a country which had fascinated
him since childhood. Most of his time in Australia was spent with the Bread Research Institute in Sydney. He
also made several extended trips to China, serving as guest lecturer at ShanDong Agricultural University, TaiAn,
and at Henan University of Technology, Zhengzhou.
Chuck always commented that the best legacy a person could leave is the knowledge he has shared. A
significant way in which his knowledge has been shared is represented by his contributions to the first and
second editions of Elseviers Encyclopedia of Food Grains.

Colin Wrigley on behalf of the Editors.

Professor Koushik Seetharaman

In August 2012, Koushik Seetharaman was appointed as the third editor of this
second edition of Elseviers Grains Encyclopedia, replacing Professor Walker.
Tragically, Koushiks many contributions to the Encyclopedia of Food Grains were
cut short by his untimely death at 48 years in June 2014.
Koushik received his BS at the Gujarat Agricultural University, Anand, Guja-
rat, India. His MS training was at Cornell University, Ithaca, NY, USA, and his
PhD research was carried out at Texas A&M University, College Station, TX, USA.
At the time of Koushiks appointment as encyclopedia editor (August 2012),
he was Associate Professor and Cereals Chair in the Department of Food Science
at the University of Guelph, Canada. Research in his laboratory spanned the
cereal-grain value chain, focusing on grain quality, processing and ingredient
interactions, and on consumer acceptability and health.
His research on grain quality at Guelph involved collaboration with cereal-breeding companies, grain
farmers of Ontario and local processors, by analyzing Ontario wheats for functionality and nutritional
attributes. Special accents were on antioxidant activity, and on fiber and phenolics contents. Interactions
with breeders and processor related to the chemistry, sensory properties and consumer acceptance of whole
grain products, with specific focus on red versus white wheats in different product matrices.
Koushik had a special interest in the behavior of starch in water-poor systems, such as dough and baked
products. His research group had demonstrated that the constituent polymers of wheat starch are a series of
biopolymers ranging from linear to branched, depending on several factors including genetics and growth
environment. Moreover, the proportion of these polymers further defines their functional properties.
Research funding had been provided by the Canadian Agricultural Adaptation Council with contributions
from the Ontario Cereal Industry Research Council, Kellogg, Kraft, Kraft Canada Mill, Dow AgroSciences, C&M
Seeds, Grain Farmers of Ontario and Brabender GmbH.
Several months before his death, Koushik had been appointed as an Associate Professor to the Department
of Food Science and Nutrition at the University of Minnesota, St Paul, MN, USA, there taking the position of
General Mills Cereal Chair. In that capacity, Koushik had again initiated collaborations with breeders and
geneticists, millers and processors, with a focus on what product quality means for the consumer. His research
focused on interactions of grain biopolymers starch and gluten in particular and the impact on processing,
product attributes and consumer health.
 Tributes to Former Co-Editors xxi

His group included three postdoctoral research associates, five doctoral students and one MS student. They
were collaborating extensively with research groups around the world including University of Milan, Italy;
University of Copenhagen, Denmark; Lille University, France; ONIRIS, France; Tamilnadu Agricultural Uni-
versity, India; Iowa State University, USA; African Rice Research Institute, Benin; University of Guelph, Canada;
and Memorial University, Newfoundland, Canada.
Koushiks research background and his collaborations placed him well to contribute to the preparation of
the grains encyclopedia, both by his our writing and via his contacts to engage world-authority contributors.
The grain-science community world-wide was shocked by Koushiks sudden passing. Koushik left a wife
Debra Freedman and an eight-year-old son, Samuel.
Koushiks contribution to the encyclopedia was such that Elsevier decided that his name should remain as a
co-editor even though he did not live to see the end result of his contributions.

Colin Wrigley on behalf of the Editors.

This page intentionally left blank
EDITORS BIOGRAPHY

Colin W. Wrigleys 55 years in grain-science research have earned him international recognition in
the form of several international and Australian research awards. His work is described in about
600 research publications, including several patents, a series of eight books on Australian cereal
varieties, and many edited books. He was Editor-in-Chief of the first edition of the Encyclopedia of
Grain Science (2004).
His research interests have centered on the characterization of cereal-grain proteins in relation to
processing quality. This has involved developing new methods of protein fractionation, including
gel isoelectric focusing and its 2D combination with gel electrophoresis, leading into proteomic
mapping. Other diagnostic methods developed relate to the evaluation of grain quality in wheat
and barley, such as better methods for variety identification and for characterizing quality in starch
and sprouted grain (as co-patentee of the Rapid ViscoAnalyser). Research involvement has also
included elucidation of grain-quality variation due to environmental factors (heat stress, fertilizer
use, CO2 levels and storage conditions).
In 2009, Wrigley was appointed a Member of the Order of Australia (AM) for service to primary
industry, particularly to grain science as a researcher, and to the development of methods for
improving wheat quality.
He is currently an Honorary Professor at the University of Queensland, Brisbane, Australia.

Harold Corke is a professor in the Food and Nutritional Sciences program at The University of
Hong Kong, and a Chutian Scholar Distinguished Foreign Professor in the Glyn O. Phillips
Hydrocolloid Research Center at HUT, Hubei University of Technology, Wuhan, China. In more
than 22 years in Hong Kong, he has had responsibility for teaching a diverse array of food science
courses, including the major courses Grain Production and Utilization, Food Safety and Quality
Management, Food Product Development, and general education courses Feeding the World, and
Food: Technology, Trade and Culture. He is author or co-author of 180 refereed journal articles,
and his 22 PhD graduates have gone on to successful careers in academia and industry around the
world. He is on the editorial boards of Journal of Food Composition and Analysis, LWT Food Science and
Technology and Journal of Cereal Science, and with Colin Wrigley and Chuck Walker was one of the
editors of the first edition of Encyclopedia of Grain Science, published in 2004. Corke is active in
consulting in grain processing and food safety in Asia and Eastern Europe.

Jon Faubion is the Charles Singleton Professor of Baking and Cereal Science in the Department of
Grain Science, Kansas State University. Over his 35 year career, he has had the sole or shared
responsibility for teaching nine different cereal or food science courses at Texas A&M University,
The University of Minnesota and Kansas State. In 2011, he received the Excellence in Teaching
Award from the American Association of Cereal Chemists International. He is a senior editor of
Cereal Chemistry, and a member of the Board of Directors of the American Association of Cereal
Chemists International and an executive editor of Journal of the Science of Food and Agriculture. Prior
to rejoining the department faculty in 2005, he directed the Applied Technology and Sensory
Science Groups for the research and development arm of The Schwan Food Company.

xxiii
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EDITORIAL ADVISORY BOARD

Jinsong Bao Bob French

Zhejiang University, Hangzhou, PR China
Frank Bekes
Director R&D FBFD Pty Ltd, Beecroft, NSW, Australia
Trust Beta
University of Manitoba, Winnipeg, MB, Canada
Stanley Cauvain
BakeTran, Witney, United Kingdom; Curtin University,
Perth, WA, Australia
Robert Cracknell
Crackers Consulting, Mount Eliza, VIC, Australia
Yapeng Fang
Hubei University of Technology, Wuhan, PR China
Jon Faubion
Kansas State University, Manhattan, KS, USA
Department of Agriculture and Food, Western Australia,
Merredin, WA, Australia
Robert Graybosch
Agricultural Research Service, USDA, Lincoln, NE, USA
Robert Henry
University of Queensland, Brisbane, QLD, Australia
Barry V. McCleary
Megazyme International Ireland, Bray, Wicklow, Ireland
Lindsay OBrien
University of Sydney, Narrabri, NSW, Australia
Paul Scott
USDA-ARS Corn Insects and Crop Genetics Research
Unit, Ames, IA, USA
John Reginald Nuttall Taylor
University of Pretoria, Pretoria, South Africa

xxv
This page intentionally left blank
CONTRIBUTORS

E Akhunov BJ Blaney
Kansas State University, Manhattan, KS, USA The University of Queensland, St Lucia, Australia
PC Annis JE Bock
CSIRO Entomology, Canberra, ACT, Australia University of Guelph, Guelph, ON, Canada
GA Annor F Bonomi
University of Ghana, Accra, Ghana University of Milan, Milan, Italy
CA Atkins AR Bridges
The University of Western Australia, Crawley, WA, AACC International, St Paul, MN, USA
Australia
H Brinch-Pedersen
W Atwell Aarhus University, Slagelse, Denmark
Bill Atwell Consulting, Champlin, MN, USA
G Bultosa
M Baga Haramaya University, Dire Dawa, Ethiopia; Botswana
University of Saskatchewan, Saskatoon, SK, Canada College of Agriculture, Gaborone, Botswana
JS Bao W Bushuk
Zhejiang University, Hangzhou, PR China University of Manitoba, Winnipeg, MB, Canada
VJ Barthet YZ Cai
Canadian Grain Commission, Winnipeg, MB, Canada China National Seed Group, Beijing, PR China;
SA Barton Huazhong Agricultural University, Wuhan, PR China;
Oxford Agricultural Trials Ltd, Oxford, UK The University of Hong Kong, Hong Kong, PR China;
Hubei University of Technology, Wuhan, PR China
IL Batey
Sunset Cereal Services, Meadowbank, NSW, Australia EF Caldwell
AACC International, Roseville, MN, USA
F Bekes
FBFD PTY LTD, Beecroft, NSW, Australia KJ Carpenter
University of California, Berkeley, CA, USA
AR Belcher
Oregon State University, Corvallis, OR, USA JC Catullo
INTA, Buenos Aires, Argentina
JN BeMiller
Purdue University, West Lafayette, IN, USA SP Cauvain
BakeTran, Witney, OX, UK
T Beta
University of Manitoba, Winnipeg, Manitoba, Canada S Ceccarelli
Via delle Begonie 2, Ascoli Piceno, Italy
JL Black
John L Black Consulting, Warrimoo, NSW, Australia GK Chandi
University of Guelph, Guelph, ON, Canada
MH Blanco
(Retired) Plant Breeder and Geneticist, Encinitas, N Chaudhary
CA, USA G. J. University of Science and Technology, Hisar, India

xxvii
xxviii Contributors

RN Chibbar JE Dexter
University of Saskatchewan, Saskatoon, SK, Canada Canadian Grain Commission, Winnipeg, MB, Canada
M Chisi B Dobraszczyk
Golden Valley Research Station, Chisamba, Zambia Reading Science Centre, Reading, UK
S Cloutier D Domingues
Agriculture and Agri-Food Canada, Ottawa, ON, General Mills Inc., Minneapolis, MN, USA
Canada
C Dowd
LS Collado General Mills Inc., Minneapolis, MN, USA
The University of the Philippines Los Banos, Laguna,
LM Duizer
Philippines
University of Guelph, Guelph, ON, Canada
NC Collins
RW Duncan
School of Agriculture Food and Wine, The University of
University of Manitoba, Winnipeg, MB, Canada
Adelaide, Glen Osmond, SA, Australia
KG Duodu
H Corke
University of Pretoria, Pretoria, South Africa
The University of Hong Kong, Hong Kong, Pokfulam,
PR China; Hubei University of Technology, Wuhan, SR Eckhoff
PR China (Retired) University of Illinois, Urbana, IL, USA
RL Cracknell M Edney
Crackers Consulting, Mount Eliza, VIC, Australia Canadian Grain Commission, Winnipeg, MB, Canada
GB Crosbie M Emery
Crosbie Grain Quality Consulting, East Fremantle, WA, Iowa State University, Ames, IA, USA
Australia
W Erskine
JS Croser University of Western Australia, Crawley, WA, Australia
University of Western Australia, Crawley, WA, Australia
WD Eustace
A Cruickshank (Retired) Kansas State University, Manhattan, KS, USA
DAFF Queensland, Warwick, QLD, Australia
A Evans
A Cuesta-Marcos Tate and Lyle, Hoffman Estates, IL, USA
Oregon State University, Corvallis, OR, USA
E Evans
MG DEgidio The Tassie Beer Dr, Lindisfarne, TAS, Australia
Unita di ricerca per la Valorizzazione Qualitativa dei
JD Faris
Cereali, Rome, Italy
USDA-ARS Cereal Crops Research Unit, Fargo, ND,
D Dan Ramdath USA
Agriculture and Agri-Food Canada, Guelph, ON, Canada
J Faubion
P Dangi Kansas State University, Manhattan, KS, USA
G. J. University of Science and Technology, Hisar, India
AM Fehily
JK Daun Tinuviel Software, Warrington, UK
(Deceased)
T Filichkin
C David Oregon State University, Corvallis, OR, USA
ISARA Lyon, Lyon, France
GB Fincher
L Day Australian Research Council Centre of Excellence in Plant
AgResearch Ltd., Palmerston North, New Zealand Cell Walls, University of Adelaide, Glen Osmond, SA,
Australia
D Deneke
South Dakota State University, Brookings, SD, USA SP Fisk
Oregon State University, Corvallis, OR, USA
R DePauw
Agriculture and AgriFood Canada, Swift Current, SK, MT Fletcher
Canada The University of Queensland, St Lucia, Australia
 Contributors xxix

RJ Fletcher A Gunaratne
University of Queensland, Gatton, QLD, Australia The University of Hong Kong, Hong Kong, PR China
F Fleurat-Lessard YN Guragain
INRA, Mycology and Food Safety Research Unit, Kansas State University, Manhattan, KS, USA
Villenave dOrnon, France
RCE Guy
JA Fox Campden and Chorleywood Food Research Association,
Kansas State University, Manhattan, KS, USA Chipping Campden, UK
RJ French C Hall III
The University of Western Australia, Crawley, WA, North Dakota State University, Fargo, ND, USA
Australia
LM Hall
B Friebe University of Alberta, Edmonton, AB, Canada
Kansas State University, Manhattan, KS, USA
EG Hammond
BJ Furman Iowa State University, Ames, IA, USA
San Jose State University, San Jose, CA, USA
R Haraszi
M Gangola Campden BRI, Gloucestershire, UK
University of Saskatchewan, Saskatoon, SK, Canada
J Harnack
P Gelinas University of Minnesota, Minneapolis, MN, USA
Food Research and Development Centre, Agriculture and
D Hauge
Agri-Food Canada, Saint-Hyacinthe, QC, Canada
Grains for Health Foundation, St. Louis Park, MN, USA
SM Ghazani
AMR Hayes
University of Guelph, Guelph, ON, Canada
St. Catherine University, Lafayette, MN, USA
MC Gianibelli
PM Hayes
(Retired)
Oregon State University, Corvallis, OR, USA
A Gianinetti
L Helgerson
Consiglio per la ricerca in agricoltura e lanalisi
Oregon State University, Corvallis, OR, USA
delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Italy RJ Henry
University of Queensland, Brisbane, QLD, Australia
BS Gill
Kansas State University, Manhattan, KS, USA D Herb
Oregon State University, Corvallis, OR, USA
JS Godber
LSU Agricultural Center, Baton Rouge, LA, USA I Herencia
Universidad Politecnica de Madrid, Madrid, Spain
A Goldstein
University of Minnesota, St. Paul, MN, USA E Hernandez
Texas A&M University, College Station, TX, USA
R Graebner
Oregon State University, Corvallis, OR, USA KB Hobson
New South Wales Department of Primary Industries,
S Graham-Acquaah
Tamworth, NSW, Australia
Africa Rice Center, Cotonou, Republic of Benin
M Hochhalter
RA Graybosch
North Dakota State University, Fargo, ND, USA
USDA-ARS, University of Nebraska Lincoln, Lincoln,
NE, USA RD Horsley
North Dakota State University, Fargo, ND, USA
S Grundas
Bohdan Dobrzanski Institute of Agrophysics of the Polish W Huang
Academy of Sciences, Lublin, Poland Jiangsu Academy of Agricultural Sciences, Nanjing,
PR China; Jiangnan University, Wuxi, Jiangsu, PR China
TJ Gulya
USDA-ARS, Northern Crop Science Laboratory, Fargo, Y Huang
ND, USA China Agricultural University, Beijing, PR China
xxx Contributors

V Humphrey-Taylor P Koehler
(Retired) New Zealand Institute for Crop Food Research Deutsche Forschungsanstalt fur Lebensmittelchemie,
Limited, Christchurch, New Zealand Leibniz Institut, Freising, Germany
T Hymowitz R Korczak
University of Illinois at Champaign-Urbana, Urbana, University of Minnesota, St. Paul, MN, USA
IL, USA
SG Krattinger
S Iametti University of Zurich, Zurich, Switzerland
University of Milan, Milan, Italy
L Krishnamurthy
C Isaak International Crops Research Institute for the Semi-Arid
University of Manitoba, Winnipeg, MB, Canada Tropics, Patancheru, Andhra Pradesh, India
MS Izydorczyk S Kumar
Canadian Grain Commission, Winnipeg, MB, Canada International Center for Agricultural Research in the Dry
Areas (ICARDA), Rabat, Morocco
P Jacques
Tufts University, Boston, MA, USA OR Larroque
CSIRO Agriculture Flagship, Canberra, Australia
S Jaiswal
University of Saskatchewan, Saskatoon, SK, Canada JW Lawton
J Jane (Retired) ARS-USDA
Iowa State University, Ames, IA, USA EA Lee
LA Johnson University of Guelph, Guelph, ON, Canada
Iowa State University, Ames, IA, USA EC Leonard
JM Jones AgriKnowHow, Maitland, SA, Australia
St. Catherine University, Arden Hills, MN, USA JS Li
A Juhasz China Agricultural University, Beijing, PR China
Centre for Agricultural Research of the Hungarian WD Li
Academy of Sciences, Martonvasar, Hungary Shanxi University, Taiyuan, PR China
BO Juliano Z Li
Philippine Rice Research Institute Los Banos, Laguna, CSIRO Food Futures National Research Flagship,
Philippines Canberra, ACT, Australia; CSIRO Plant Industry,
RS Kadan Canberra, ACT, Australia
(Deceased)
R Lindberg
GP Kadkol Minneapolis Heart Institute Foundation, Minneapolis,
NSW Department of Primary Industries, Calala, NSW, MN, USA
Australia
MG Lindhauer
LW Kannenberg Max Rubner-Institut (MRI), Detmold, Germany
University of Guelph, Guelph, ON, Canada
K Liu
TN Khan Agricultural Research Service, US Department of
University of Western Australia, Crawley, WA, Australia Agriculture, Aberdeen, ID, USA

BS Khatkar Q Liu
G. J. University of Science and Technology, Hisar, India Agriculture and Agri-Food Canada, Guelph, ON, Canada

JG Kling S Lu
Oregon State University, Corvallis, OR, USA National Chung Hsing University, Taichung, Taiwan
EJ Knights OM Lukow
New South Wales Department of Primary Industries, Agriculture and Agri-Food Canada, Winnipeg, MB,
Tamworth, NSW, Australia Canada
 Contributors xxxi

BL Ma NJ Mendham
Agriculture and Agri-Food Canada, Ottawa, ON, University of Tasmania, Hobart, TAS, Australia
Canada
R Menon
C-Y Ma The General Mills Bell Institute of Health and Nutrition,
The University of Hong Kong, Hong Kong, PR China Minneapolis, MN, USA
A MacLeod TE Michaels
Canadian Grain Commission, Winnipeg, MB, Canada University of Minnesota, St. Paul, MN, USA
L MacLeod DM Miskelly
Dairy Innovation Australia, Werribee, VIC, Australia Westcott Consultants P/L, Goulburn, NSW, Australia
CK Madsen
CS Moffat
Aarhus University, Slagelse, Denmark
Curtin University, Bentley, WA, Australia
RJ Mailer
ES Monyo
Australian Oils Research, Lambton, NSW, Australia
ICRISAT, Nairobi, Kenya
JT Manful
Africa Rice Center, Cotonou, Republic of Benin TR Moore
AIB International, Manhattan, KS, USA
AG Marangoni
University of Guelph, Guelph, ON, Canada MK Morell
International Rice Research Institute, Los Banos,
DJ Mares Philippines
University of Adelaide, Glen Osmond, SA, Australia
CF Morris
L Marquart USDA-ARS Western Wheat Quality Laboratory,
University of Minnesota, St. Paul, MN, USA; Grains for Pullman, WA, USA
Health Foundation, St. Louis Park, MN, USA
LA Morrison
A Marti
Oregon State University, Corvallis, OR, USA
Universita degli Studi di Milano, Milan, Italy
B Maschoff WK Mousa
Grains for Health Foundation, St. Louis Park, MN, USA University of Guelph, Guelph, ON, Canada; Mansoura
University, Mansoura, Egypt
A Mathiowetz
University of Minnesota, St. Paul, MN, USA; Rochester, K Mrva
MN, USA University of Adelaide, Glen Osmond, SA, Australia

JJ Mayko L Munck
Canola Council of Canada, Mundare, AB, Canada University of Copenhagen, Frederiksberg C, Denmark
CM McDonough R Munns
Texas A&M University, College Station, TX, USA CSIRO Agriculture, Canberra, ACT, Australia; The
University of Western Australia, Crawley, WA, Australia
RA McIntosh
The University of Sydney, Cobbitty, NSW, Australia PA Murphy
Iowa State University, Ames, IA, USA
JD McKeehen
Cereal Partners Worldwide, Minneapolis, MN, USA ED Nafziger
J McLean University of Illinois, Urbana, IL, USA
The University of Newcastle, Callaghan, NSW, Australia SC Nelson
PBE McVetty AACC International (AACCI), St. Paul, MN, USA
University of Manitoba, Winnipeg, MB, Canada T Nleya
B Meints South Dakota State University, Brookings, SD, USA
Oregon State University, Corvallis, OR, USA
L OBrien
A Meldrum The University of Sydney, Narrabri, NSW, Australia;
Pulse Australia, Perth, WA, Australia Solheimar Pty Ltd, Narrabri, NSW, Australia
xxxii Contributors

DK OToole I Rajcan
City University of Hong Kong, Hong Kong, PR China University of Guelph, Guelph, ON, Canada
EA Oelke L Ramsden
University of Minnesota, St. Paul, MN, USA University of Hong Kong, Hong Kong, PR China
RP Oliver KD Rausch
Curtin University, Bentley, WA, Australia University of Illinois at UrbanaChampaign, Urbana, IL,
USA
DE Ortiz
W. K. Kellogg Institute, Battle Creek, MI, USA A Regina
MA Pagani CSIRO Food Futures National Research Flagship,
Universita degli Studi di Milano, Milan, Italy Canberra, ACT, Australia; CSIRO Plant Industry,
Canberra, ACT, Australia
TS Palav
Rich Products Corporation, Buffalo, NY, USA MN Riaz
Texas A&M University, College Station, TX, USA
L Pallas
Yanco Agricultural Institute, Yanco, NSW, Australia D Richard-Molard
(Retired) INRA, Nantes, France
GH Palmer
Heriot-Watt University, Edinburgh, UK F Rizza
Consiglio per la ricerca in agricoltura e lanalisi
RG Palmer delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Iowa State University, Ames, IA, USA Italy
RF Park MJ Robertson
The University of Sydney, Narellan, NSW, Australia CSIRO Sustainable Ecosystems, St. Lucia, QLD,
GM Paulsen Australia
Kansas State University, Manhattan, KS, USA LW Rooney
LC Paulson Texas A&M University, College Station, TX, USA
The Bama Companies, Tulsa, OK, USA WL Rooney
DS Petterson Texas A&M University, College Station, TX, USA
Department of Agriculture, Nedlands, WA, Australia CM Rosell
RA Porter Institute of Agrochemistry and Food Technology (IATA-
University of Minnesota-NCROC, Grand Rapids, CSIC), Paterna, Valencia, Spain
MN, USA AS Ross
L Privalle Oregon State University, Corvallis, OR, USA
Bayer CropScience, Research Triangle Park, Durham, SJ Roy
NC, USA School of Agriculture Food and Wine, The University of
KV Probst Adelaide, Glen Osmond, SA, Australia
Kansas State University, Manhattan, KS, USA; Grain HD Sapirstein
Processing Corporation, Muscatine, IA, USA University of Manitoba, Winnipeg, MB, Canada
KJ Quail AK Sarkar
Grain Growers Limited, North Ryde, NSW, Australia Canadian International Grains Institute, Winnipeg, MB,
RCN Rachaputi Canada
University of Queensland, Kingaroy, QLD, Australia
A Sarker
H Rahman International Center for Agricultural Research in the Dry
University of Alberta, Edmonton, AB, Canada Areas (ICARDA), New Delhi, India
S Rahman SK Sathe
Monash University Malaysia, Bandar Sunway, Malaysia Florida State University, Tallahassee, FL, USA
MN Raizada SF Schakel
University of Guelph, Guelph, ON, Canada University of Minnesota, Minneapolis, MN, USA
 Contributors xxxiii

KA Scherf VA Solah
Deutsche Forschungsanstalt fur Lebensmittelchemie, Curtin University, Perth, WA, Australia
Leibniz Institut, Freising, Germany
AM Stanca
MP Scott University of Modena and Reggio Emilia, Reggio Emilia,
Iowa State University, Ames, IA, USA Italy
K Seetharaman FL Stoddard
(Deceased) University of Helsinki, Helsinki, Finland
GJ Seiler K-C Tan
USDA-ARS, Northern Crop Science Laboratory, Fargo, Curtin University, Bentley, WA, Australia
ND, USA
AS Tatham
SO Serna-Saldivar Cardiff Metropolitan University, Cardiff, UK
Centro de Biotecnologa FEMSA, Escuela de Ingeniera y
JRN Taylor
Ciencias, Monterrey, Mexico
University of Pretoria, Hatfield, South Africa
S Shea Miller
V Terzi
Eastern Cereals and Oilseeds Research Centre,
Consiglio per la ricerca in agricoltura e lanalisi
Agriculture and Agri-Food Canada, Ottawa, ON,
delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Canada
Italy
DB Sheats
K Tiefenbacher
St. Catherine University, St. Paul, MN, USA
(Retired)
PR Shewry
VK Tiwari
Rothamsted Research, Harpenden, UK; University of
Kansas State University, Manhattan, KS, USA
Reading, Reading, UK
SM Tosh
RD Shillito
Agriculture and Agri-Food Canada, Guelph, ON, Canada
Bayer CropScience, Durham, NC, USA
P Tosi
JP Shroyer
University of Reading, Reading, UK
Kansas State University, Manhattan, KS, USA
LC Trugo
KJ Shroyer
Federal University of Rio de Janeiro, Rio de Janeiro, Brazil
Kansas State University, Manhattan, KS, USA
SE Ulrich
Qingyao Shu
Washington State University, Pullman, WA, USA
Zhejiang University, Hangzhou, PR China
EH Unger
KHM Siddique
Eagle Rock Solutions, Inc., Idaho Falls, ID, USA
University of Western Australia, Crawley, WA, Australia
PV Vadlani
J Singla
Kansas State University, Manhattan, KS, USA
University of Zurich, Zurich, Switzerland
SA Valencia-Chamorro
M Sissons
Escuela Politecnica Nacional, Quito, Ecuador
NSW Department of Primary Industries, Calala, NSW,
Australia; Tamworth Agricultural Institute, Tamworth, V Vamadevan
NSW, Australia University of Minnesota, St. Paul, MN, USA; Agriculture
and Agri-Food Canada, Guelph, ON, Canada
JH Skerritt
University of Queensland, St Lucia, QLD, Australia N Van Heel
University of Minnesota, Minneapolis, MN, USA
DJ Skylas
Grain Growers Limited, North Ryde, NSW, A Vandenberg
Australia University of Saskatchewan, Saskatoon, SK, Canada
DL Smith SK Vasal
McGill University, Ste-Anne-de-Bellevue, QC, Canada BISA, Punjab, India
xxxiv Contributors

R Velicogna BJB Wood

Archer Daniels Midland, Montreal, QC, Canada Strathclyde University, Glasgow, UK
D von Baer G Wright
Seeds Baer, Temuco, Chile Peanut Company of Australia, Kingaroy, QLD, Australia

E von Baer CW Wrigley

CAMPEX Semillas Baer, Temuco, Chile
CE Walker
(Deceased)
SB Walker
University of Guelph, Guelph, ON, Canada
FL Walley
University of Saskatchewan, Saskatoon, SK, Canada
D Wang
Huazhong Agricultural University, Wuhan, PR China
T Wang
Iowa State University, Ames, IA, USA
RD Waniska
Texas A&M University, College Station, TX, USA
LL Ward
Kansas State University, Manhattan, KS, USA
H Wieser
Deutsche Forschungsanstalt fur Lebensmittelchemie,
Leibniz Institut, Freising, Germany
RM Williams
CBH Group, West Perth, WA, Australia
PM Williamson
(Retired) Leslie Research Centre, Toowoomba, QLD,
Australia; CBH Group, West Perth, WA, Australia
Honorary Professor, QAAFI; The University of
Queensland, Brisbane, QLD, Australia
Dianxing Wu
Zhejiang University, Hangzhou, PR China
HX Wu
Wuhan Chortle Biotechnology Company Limited,
Wuhan, PR China
J Wyatt
NIAB, Cambridge, UK
Y Yan
Huazhong University of Science and Technology, Wuhan,
PR China
H Yangcheng
Iowa State University, Ames, IA, USA
V Zevallos
Johannes Gutenberg Mainz, Mainz, Germany; University
Medical Center, Mainz, Germany
XM Zhou
McGill University, Ste-Anne-de-Bellevue, QC, Canada
PK Zwer
South Australian Research and Development Institute
(SARDI), Urrbrae, SA, Australia
S Zydenbos
(Retired) New Zealand Institute for Crop Food Research
Limited, Christchurch, New Zealand
 THE WORLD OF FOOD GRAINS
The Basics
Contents
The Grains that Feed the World
Course Structures: Based on EFG Articles
The Grain Crops: An Overview
The Grain Chain: The Route from Genes to Grain-Based Products
Taxonomic Classification of Grain Species
Grain: Morphology of Internal Structure
Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
The Grains that Feed the World
CW Wrigley, University of Queensland, Brisbane, QLD, Australia
H Corke, University of Hong Kong, Hong Kong, Pokfulam, China
J Faubion, Kansas State University, Manhattan, KS, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Grains are the source of half of our food and protein
consumption.
Current world production for all grains equals 1 kg per
person per day.
Grains also contribute to the wider range of foods via
animals.
Major media claims about grains and health are presented
in this article.
Some myths about grain-based foods are busted.
Some claims are supported, at least in part.
Learning Objectives
To understand the significance of grains in feeding the
growing human population
To adopt a critical attitude to production and nutritional
information about grains presented in the media and to
know how to seek and research authentic information
To be motivated to pursue further topics in depth, starting
with investigating relevant articles in the Encyclopedia of
Food Grains and going back where necessary to primary
scientific research articles.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00003-6
Introduction
Grains are responsible directly or indirectly for most of the
food we eat. The big three cereal grains (rice, wheat, and
maize) have roles as staple foods (major daily sources of
carbohydrate-based energy) and for processing into animal
feeds and other diverse uses such as fuel bioethanol. There
has been an explosive growth in meat consumption globally,
as more and more people come out of poverty and enter into a
growing middle class. This is most apparent in China, where
demand for meat and other animal products (eggs and milk)
drives the production of maize domestically and production of
soy beans in major exporting countries (the United States,
Argentina, and Brazil).
A billion people worldwide are obese; a third of the food
produced in industrialized countries is thrown away uneaten.
In this exuberant excess of food production lie many serious
problems a billion people worldwide are chronically mal-
nourished, food production competes with industry and
domestic usage for scarce resources of energy and water,
human populations continue to increase, and food production
impacts on the environment in ways that may not be sustain-
able. People care about what they eat, about their health, about
longevity, and about fitness.
In the news media, nutrition sells; there is thus a constant
output of information on topics such as gluten-free diets,
1
2 THE BASICS | The Grains that Feed the World

obesity, genetically modified organisms (GMOs), and organic

agriculture. Unfortunately, much of the popular media report-
ing on grains and health is wrong, sometimes the result of
journalistic carelessness or ignorance, sometimes from
demand for an ideological spin on knowledge, and sometimes
out of a crass calculation that hysteria sells better than truth.
This encyclopedia sets out to redress the balance and rectify
these popular misconceptions. What do you expect from an
encyclopedia? Obviously, everything you want to know about
the general topic. Just as importantly, you expect correct infor-
mation, provided by world-recognized authorities, knowing
that all facts have been checked. These expectations are fulfilled
in this encyclopedia, in its four print volumes, in its hundreds
of articles, in its broad-ranging website, and in its extension as
part of Elseviers Food Science Reference Module, as a general
source of advice to the food industry worldwide.
Often, the expectation of an encyclopedia is to check
whether what-I-have-heard is actually so. This introductory
article explores some of those what-I-have-heard sayings, listed
in groups of dot points, with commentary, plus referral to
relevant encyclopedia articles. The authors of the encyclopedia
articles are recognized as world authorities in their respective
fields. The articles have all undergone peer review by members
of the editorial advisory board and the editors, thereby ensur-
ing that the information is correct and free from the many
myths, misconceptions, and misinformation that have been
spread by nefarious advertising and unbridled websites.

Grains and Our Future

Climate change will lead to less grain production. Not

necessarily!
We have become so dependent upon eating cereal grains
(grass seeds) that it has prompted at least one author to say
that we have become canaries. Yes, we are dependent on
grains, but canaries, no!
Cereal grains literally stand between humankind and
starvation. Maybe.
Humankind will run out of food in the foreseeable future.
Probably not.
What ultimately is the source of our food? How vulnerable
are we?
Our Dependence on Grains as Food
The 1960s were a time of great pessimism about the future of
humanity and the possibility of massive famines due to popu-
lation increases, especially in Asia and Africa, and the failure of
national agricultural to produce enough to feed these growing
populations. The Green Revolution was spearheaded by the
research of Dr. Norman Borlaug (Figure 1) on dwarf wheat at
CIMMYT in Mexico and was complemented by similar work
on dwarf rice at International Rice Research Institute in the
Philippines. These advances led to rapid increases in grain
production. India went from being a chronic major importer
of wheat to avert famine to being a significant exporter of grain.
The relative abundance in grain production today, the
power of biotechnology, and the excesses in consumption we
often see around us (wastage of food and increasing meat
consumption) may make us forget how tenuous is our ability
to produce the grains we need year after year. Humankind
has certainly become dependent upon eating cereal grains.
Figure 1 Norman Borlaug was awarded the Nobel Peace Prize in
1970 for his work to end world hunger by creating new strains
of drought-resistant wheat. This statue was placed in the US Capitol
Buildings National Statuary Hall in 2014 to celebrate the 100th
anniversary of Borlaugs birth.
This dependence is inevitable, given our vast population and
the contribution of grains to food and feed production. Grains
provide over half our protein and energy needs. But it is hardly
justified to claim that, as a result, we have become canaries.
A recurring theme of this encyclopedia is the great diversity of
ways in which we prepare grains in our diet not only as our
daily bread (Bible, Matthew 4:4) and not only as the staple
food of boiled rice. A complete volume is devoted to the great
diversity of grain-based products and their processing.
What Ultimately Is the Source of Our Food on Earth?
This question was addressed in 1648 by a Belgian scientist, Jan
Baptist van Helmont, with respect to plants in general. He
planted a tree in a pot of soil. He first weighed the soil (after
drying it) and also the small tree. For 5 years, he added water
only to the pot. He then weighed the remaining soil and the
tree, which had grown considerably. There was negligible
weight loss from the soil, so he concluded that all the weight
gain of the tree had come from the water that he added: thus,
wood is made from water! (Figure 2).

Today, we know that Jan did not control two important
factors: the sun and the air (CO2), which together produce all
plant life via photosynthesis according to the following reactions:
CO2 H2 O light ! O2 glucose a simple sugar
Glucose ! carbohydrates, lignin, and cellulose
plant components
Jan Baptist lived in an age of the Aristotelian belief in four
elements: water, earth, air, and fire. Even considering these
elements, he took into account the first two, but neglected air
(CO2) and fire (sunlight).
Thus, all our food comes from these sources sunlight,
CO2, and water either directly in the cases of grains,
vegetables, and fruit or indirectly in the cases of animals who
eat the grain and plants. In addition, the soil does contribute
In 1648, a scientist
provided water only
to a small tree
... for 5 years
Thus, he concluded
that trees are made from
water! He neglected the
roles of light and CO2
Figure 2 The experiment of Belgian scientist, Jan Baptist van Helmont,
seeking to determine what plants are made from.
Cereal production, utilization and stocks
Million tonnes
2600
2400
2200
2000
1800
2004/05 2006/07 2008/09
Production (left axis)
Stocks (right axis)
Figure 3 The past decade of production and utilization of cereal grains. Source: FAOSTAT.
THE BASICS | The Grains that Feed the World 3
the wide range of elements described later in this article, espe-
cially nitrogen for protein synthesis. When grain is harvested,
these elements are removed from the field, and they must
ultimately be replaced by some form of fertilizer.
However, the other essentials sunlight, CO2, and water are
readily available for free on an ongoing basis. Of course, that
might not be claimed for water in many dry climates. The free
claim can certainly be made for CO2 given the rising levels of
this component of our atmosphere. In fact, the sequestration of
CO2 by plants is one of the many solutions for mitigating the
rising levels of CO2. There needs to be a realization of the
essential nature of CO2 as the source of all plant life and thus
of all our food, thereby to reverse the bad press attributed to CO2.
Will We Run Out of Food in the Foreseeable Future?
Our reliance on the grains of cereal plants (especially wheat,
rice, and corn) is so great that they supply nearly 60% of our
food energy and 50% of the protein consumed on Earth. Thus,
grains really do stand between us and starvation. Grains offer
the great advantages that they can be stored safely for long
periods and thus transported far from their growth sites. Grains
are adaptable with respect to the wide range of foods made
from them. Grains are also important as animal food and for
various forms of industrial processing.
But will we run out of food in the foreseeable future? Efforts
continue to improve methods of breeding, agronomy, harve-
sting, and processing, showing good signs that grain produc-
tion and quality will continue to increase, providing for a
world of hungry people. The production and yield of wheat
have tripled in the past 40 years, as explained in the Wheat
Overview article (The Cereal Grains: Wheat: An Overview of
the Grain That Provides Our Daily Bread). Furthermore,
Figure 3 shows how there is an ongoing increase in the
Million tonnes
800
600
400
200
0
2010/11 2012/13 2014/15
fcast
Utilization (left axis)
4 THE BASICS | The Grains that Feed the World
Earths atmospheric CO2 concentration and temperature
390
370
350
ppm 330
CO2
310 CO2
290
270
250
1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 2000
Year
Figure 4 Increasing temperatures and levels of carbon dioxide promise increasing grain yields due to the carbon-fertilizer effect.
worldwide production of cereal grains, also including maize,
rice, barley, and other grass-related grains. The future therefore
looks hopeful.
However, we still have great inequalities in food supply in
terms of grains. Although current world production for all
grains is equivalent to 1 kg per person per day, there is still
hunger because some of this great volume of grain serves alter-
native important nonfood purposes, such as animal feed and
industrial uses. The greatest problem, however, is that the major
sites of production are distant from the major sites of need, and
those in need lack the resources to access the excess grain.
Increasing levels of carbon dioxide (Figure 4) offer the
promise of increasing grain yields, but with growth conditions
changing, breeders and agronomists need to focus research on
adapting to a changing world scene.
Climate Change and the Future of Grain Production
Predictions of global climate change include increased levels of
carbon dioxide in the atmosphere, causing warmer tempera-
tures (especially the daily minima), changed patterns of rain-
fall, and increased frequency and severity of heat-stress
episodes, together with decreased frost frequency and rising
sea levels. These trends imply a future of fluctuations in grain
yield and quality, but overall increases in grain yield are pre-
dicted. Rising sea levels have serious implications for many
low-lying rice-growing areas in Asia.
Irrespective of arguments about causes, increases in atmo-
spheric levels in carbon dioxide have been clearly demon-
strated, rising from ancient levels of 150280 to 400 ppm
and beyond, with a predicted doubling in a century or so.
One consequence is the provision of carbon fertilizer, leading
in the future to more efficient biomass production and greater
yields for many grain species. However, the yield increases may
be largely in carbohydrate, potentially leaving reduced grain-
protein levels. The fertilizing effects of carbon dioxide are
more significant at higher temperatures, the other major effect
predicted to accompany the global climate challenge. For
wheat, the increases in temperature are also expected to affect
potential dough-forming properties. Temperatures in the range
1530  C during grain filling have been shown to provide
optimal dough strength, but grain with weaker dough quality
is produced when plants are subjected to a few days of heat
14.5
14.3
14.1
C
C 13.9
13.7
13.5
stresses of over 35  C during grain filling. However, there are
indications that some wheat genotypes are available that
provide stable grain quality despite such heat-stress condi-
tions, thereby opening up promising research directions. So,
considerable research effort is concentrated on breeding for
changed growth conditions (see articles Agronomy of Grain
Growing: Implication of Climate Changes; Sustainable Grain
Production and Utilization; Wheat: Biotrophic Pathogen Resis-
tance; Genetics of Grains: Abiotic Stress Genes and Mecha-
nisms in Wheat; The Cereal Grains: An Overview of the Family
of Cereal Grains Prominent in World Agriculture).
Growing Our Grains
Organic-grown grain is superior nutritionally to other
grain. Wrong!
GMO grain is hazardous to your health. Wrong!
Organic Growing
Organic is an adjective used with the whole range of foods and
even fibers. The term is associated with a higher price and
presumed higher value. It should also indicate products of
such quality and composition as are expected from growing
these products in the absence of chemicals, instead using nat-
ural means of farm management. Are organically grown grains,
for example, any different nutritionally from other sources of
grain? The answer is generally no. Organic grain will look the
same as conventionally grown grain and will not likely to have
more favorable nutritional content.
Organic grain is likely to have a protein content similar to
that of other grains grown under similar conditions. With
respect to chemicals, the levels of pesticides and herbicides of
organic grain may be less than those for conventionally grown
grain, but the latter are still required to have such compounds
at levels below the minimum residue levels. A report by the UK
Crop Protection Association found lower levels of pesticide
residues in organic produce when compared to conventional
but these differences are relatively small, for example, a typical
residue of 0.1 mg/kg is equivalent to a fly on a ten-ton truck.
The encyclopedias article on organic grain points out that
the great advantage of organic farming is the benefit to land
 THE BASICS | The Grains that Feed the World 5

use and the environment (see articles Agronomy of Grain Grow- on the actual composition of GM food and the actual effects
ing: Organic Grain Production and Food Processing; Sustainable of GM plant and animal agricultural practices (see articles
Grain Production and Utilization; Food Grains and the Con- Food Grains and the Consumer: Genetically Modified Grains
sumer: Labelling of Grain-Based Foods). and the Consumer; Genetics of Grains: Detection of Geneti-
cally Modified Grains; Development of Genetically Modified
Grains; The Cereal Grains: Maize: Overview; The Oilseeds:
Genetically Modified Organisms Cottonseed: Overview; Canola: Overview; Soybean:
It is well over a century since grain scientists commenced the Overview).
now usual practice of cross pollination to combine the benefits of
one parental line with those of another. Previously, plant
improvements had involved the selection of promising plants Grains, Our Health, and the Media
(by appearance) from the common mix of plants in the field.
Cross pollination greatly extended the genetic variability of popu- What can we believe? We all know that the French queen Marie
lations from which to select promising new genotypes for use in Antoinette (Figure 5) famously declared Let them eat cake
developing new improved varieties. However, this approach to when she was told that her starving subjects were clamoring for
plant breeding is limited because cross pollination is generally bread. Wrong!
restricted to plants of the same genus, perhaps even the same The saying is stuck in historical tradition because it has been
species. Thus, for example, it is not possible to source genetic a convenient precis of the aristocratic arrogance of the times
resistance to a disease from beyond the target grain species. preceding the French revolution (178999). However, the line
In recent decades, this limitation has been removed by the about eating cake was actually written in the mid-1760s when
introduction of genetic engineering techniques, permitting not Marie Antoinette was a young girl living in Austria. It appears in
only very precise inter-genus transfer of genes but also the target- a work entitled Confessions written by Jean-Jacques Rousseau.
ing of single genes or groups of genes for transfer from unrelated Furthermore, much has been lost in translation. The word
genera (which could, e.g., be as distant as a fish and a different cake is a mistranslation of the French brioche that is a pastry
plant). The improved plant or animal that carries functional with tender crumb made with generous inclusion of egg and
genes from an unrelated genus (a gene transfer that could not butter, hardly like our general concept of cake. The linkage to
happen in nature) is termed genetically modified (GM). Many Marie Antoinette has been attributed to Jean-Baptiste Alphonse
improved grain genotypes have been introduced into agricul- Karr (180890), who wrote it 50 years after her death.
tural production on a very large scale. Such improvements may How does it happen that society accepts the authenticity of
include not only agronomic and disease benefits but also a phrase or story even though it is just plain wrong? The Let
improvements in nutritional value. The major crops involved them eat cake story may be no more than a trivial error of
for large-scale production are maize, soybean, canola, and cot- history. More important are the multitude of other errors and
ton. In the case of GMO cotton especially, there has been great misconceptions in the public mind. To the extent that this
environmental benefit of reduced use of insecticides. There are misinformation affects our way of life and our personal well
GMO variations of many other grain crops, such as golden rice,
GM to contain beta-carotene, a source of vitamin A.
There has been political/philosophical/activist and con-
sumer resistance to the benefits offered by GMO technology,
such that the range of GMO grains has been restricted and the
potential benefits have not yet become available. For example,
there is thus no widespread cultivation of GMO wheat, barley,
or rice. Much of the scientific evidence presented on the poten-
tial harm of GM foods is simply spurious, generated with
disregard to normal experimental practices, and with results
predetermined by a bias on the part of the authors. In conse-
quence of consumer resistance, the spread of GMO grains has
been restricted to some countries and regions mainly to the
Americas. In Argentina, for example, many farmers grow two
crops a year wheat and GMO soy. As the soy is herbicide-
resistant, weeds are eliminated during the soy-growth phase,
and the subsequent wheat crop is harvested free of weed seeds.
Much of Europe and Africa has been far more circumspect
about permitting introduction of GM grains. There is a valid
and essential ongoing public debate to be made about the
ownership or intellectual property basis of genetic resources
that underpin GMO production. There is valid unease about
the potentially excess control that a few biotechnology compa-
nies may exert on world agriculture. There are also arguments
to be made about the environmental impacts both positive
and negative of all forms of agriculture, including that
involving GM plants and animals. But these arguments should Figure 5 The peasants of France were starving for lack of bread. Marie
be made on the basis of accurate and apolitical information Antoinette responded: No bread? Then let them eat cake. Did she?
6 THE BASICS | The Grains that Feed the World
being, there is a serious need for authentic information to be
offered and adopted.
There may be no combination of issues where this need is
more apparent than in the matter of nutrition, our diet being
closely allied to our health, quality of life, and our death, thus
prompting sayings such as the following:
What we eat today walks and talks tomorrow.
Digging our graves with our teeth.
We are what we eat.
Mens sana in corpore sano (a healthy mind in a healthy
body).
The Encyclopedia of Food Grains: A Source of
Authentic Information
This encyclopedia offers authentic information about all
aspects of food grains as they relate to us and about all
sorts of grains: chickpeas, rice, maize, barley, quinoa, soya,
plus buckwheat, and the real wheat (the last two grains are
unrelated!). The encyclopedias coverage takes us through the
various stages of the breeding of grains, their growth, produc-
tion, processing, and the retailing of the wide range of grain-
based foods. All these issues impact us as consumers, situated
as we are at the culmination of the grain chain (see article The
Basics: The Grain Chain: The Route from Genes to Grain-Based
Products). Veracity for the articles is assured due to authorship
by world authorities, followed by peer review by members of
the editorial advisory board, the editors, and external experts.
Some Popular Mythconceptions About Food Grains
In this opening article, we examine and correct some of the
myths and misconceptions (mythconceptions) about grain-
related nutrition and grain production. These errors appear in
popular books and on questionable websites and inferred in
advertising, in general gossip, and in popular concepts. Beyond
this article, more mythconceptions are addressed in detail
throughout the encyclopedia. How about the following ideas?
Bread makes you fat. Wrong!
The fat you eat goes straight to your hips. Wrong!
Eat fat-free products and you wont get fat. Wrong!
Trans fats, produced in fats from oilseeds, are bad for you.
Maybe!
Food additives are all badditives! Wrong!
New types of oilseeds have fats that are nutritionally bene-
ficial. True!
Only children get celiac disease and they can grow out of it
anyway. No and no!
A gluten-free diet will make you feel better. Wrong or not
necessarily.
A gluten-free diet is damaging to your health. Wrong!
A gluten-free diet will cause you to miss essential nutrients.
Maybe.
A gluten-free diet means that I cannot eat buckwheat.
Wrong!
Figure 6 This type of corn dolly is not made from corn (maize); they are
usually made from the stalks and heads of wheat, rye, triticale, and
barley. This apparent misnomer arises from the use of corn to signify any
grain species; this connotation is still prevalent in parts of Britain,
whereas corn is used in North America to signify maize.
At least we know that milk and coffee are gluten-free. Not
necessarily.
Corn dollies are made from corn (maize). No, not usually!
(See Figure 6.)
There are dietary imbalances in the cereal grains. Partly true!
Spinach is the best source of iron in our diet; we know that
because we grew up with Popeye the Sailor Man who ate all
his spinach. Wrong!
Wild flax growing in New Zealand differs from cultivated
flax. Correct! (See Figure 7.)
Organic-grown grain is superior nutritionally to other
grain. Wrong!
GMO grain is hazardous to your health. Wrong!
Authentic information is needed on the aforementioned topics
and on many more. Ask a question about grains and seek an
authentic answer via the hypertext features or the conventional
index of this encyclopedia. The assurance of authenticity of this
encyclopedia is that world authorities have been sought to
write every chapter, and subsequently, all articles have been
peer-reviewed. None of those involved have any commercial
interest in the topic about which they have written. Those
assurances cannot be made for the authors of many popular
books, magazine articles, or websites.
What Do Our Bodies Need and How Much?
Lets get to the facts of our nutrition. Our bodies are composed
of carbon (C), hydrogen (H), oxygen (O), nitrogen (N), cal-
cium (Ca), phosphorus (P), and the wide range of other chem-
ical elements illustrated in Figure 8. These then are the
elements that are essential in our diet. Their distribution varies
across the various foods we eat, as illustrated for breakfast time
in Figure 9. The importance of the grain-based foods is evident
as a source of an important combination of these elements. In
addition to the trace elements, there are the compounds that
we cannot make ourselves, including vitamins, essential amino
acids (forming proteins), and essential fatty acids.
This information is thus the starting point for good nutri-
tion. Beyond these essential components is the matter of how

Figure 7 (a) This is flax. It is wild flax growing in New Zealand, a form
of lily (Phormium tenax and Phormium colensoi). It is used by the
Maoris to make fabrics. (b) This is flax. It is seed from cultivated flax
(Linum usitatissimum). This flax or linseed is valuable for its oil content.
In addition, the leaves are used to produce linen. The genus name
was adapted for the floor covering linoleum (informally abbreviated to
lino), which is made from solidified linseed oil (linoxyn). Flax seed is
reputed to assist in lowering serum cholesterol levels.
much of them and in what proportions. This topic has been
addressed in many countries by the provision of nutritional
guidelines, such as those in Figures 10 and 11. They empha-
size the quantitative importance of the grain-based foods in
our diets, either as the base of the pyramid or as the largest
sector of the dinner plate. In fact, the major cereal grains
(wheat, maize, rice, barley, sorghum, oats, rye, and millet)
provide about half the protein needs of mankind and over
half of our food energy consumption. Additional contribu-
tions are made by the noncereal grains, including soy, canola,
sunflower, and legumes. Worldwide, annual grain production
exceeds 2.5 billion tonnes. This equates to about 1 kg of grain
person per day for the whole worlds population! Why is
there hunger anywhere?
THE BASICS | The Grains that Feed the World 7
Finally, there is the important factor of how much food we
should consume. While an excess of food consumption is a
critical health factor in many countries, elsewhere, hunger is
still present. How can this contrast exist when grain production
is so great?
Correcting Misinformation
Overlaying the elementary recommendations coming from
Figures 8 and 9, there are many, many points of discussion,
various arguments, and gross issues of misinformation about
our nutrition. The points listed in the preceding text are cov-
ered in the succeeding text.
Grains, Fats, and Weight Loss
The fat you eat goes straight to your hips. Wrong!
Eat fat-free products and you wont get fat. Wrong!
Bread makes you fat. Wrong!
Food additives are all badditives! Wrong!
Trans fats, produced in fats from oilseeds, are bad for you.
Maybe!
New types of oilseeds have fats that are nutritionally extra
beneficial. True!
The Fat We Eat: Where Does It Go?
All the food we eat is broken down to simpler components in
our stomach and intestine before it is absorbed. Starch is
hydrolyzed and absorbed as glucose. Fats are absorbed as the
fatty acids. These simpler compounds are metabolized to
acetyl coenzyme A, which in turn is digested via the Krebs
cycle (Figure 12) to carbon dioxide and compounds that
store the resulting energy. Subsequently, these metabolites
provide energy to our muscles and replace almost all of the
material in our bodies in regular turnover activities. Some
excess will be laid down as fat deposits (maybe in the hips),
but these fat deposits come indirectly from any of the compo-
nents in our diet, not only from fats. Thus, eating fat-free
foods does not necessarily relate to weight loss or gain any
more than any other food source (see article Food Grains and
the Consumer: Grains and Health: Misinformation and
Misconceptions).
Bread as a Food
Bread provides good proportions of protein, starch, fiber, and
fat. But watch what goes onto the bread as a spread or as
sandwich filling!
Flour and ground grain are obviously the main ingredients
in bread and baked goods generally. In addition, there are
various ingredients (some sugar, salt, and fat, plus rising agents
such as yeast and baking powder), including some ingredients
that may be classed as additives. The added ingredients have
the functions of enhancing the quality and nutritional value of
the baked foods. But they certainly cannot be labeled as
8 THE BASICS | The Grains that Feed the World

OTHERS
ABUNDANT ELEMENTS

1.5%
PHOSPHORUS RARE
SULFUR
1.0%
POTASSIUM
SODIUM
CHLORINE
CALCIUM
MAGNESIUM
1.5% VERY RARE
IRON
FLUORINE
NITROGEN SILICON

3.0% ZINC
STRONTIUM
RUBIDIUM
LEAD
HYDROGEN
MANGANESE
10% COPPER
ULTRA RARE
ALUMINUM
CARBON CADMIUM
TIN
18% BARIUM
MERCURY
SELENIUM
IODINE
OXYGEN
MOLYBDENUM
65% NICKEL
BORON
CHROMIUM
ARSENIC
COBALT
VANADIUM
= MINERALS

Figure 8 The proportions of the chemical elements that make up our bodies. Reproduced from Bunpei Y (2012) Wonderful Life with the Elements. San
Francisco, CA: No Starch Press.

badditives. All such ingredients must be approved by the
respective government agencies (see articles Food Grains and
the Consumer: Fortification of Grain-Based Foods; Wheat-
Based Foods: Breads; Cookies, Biscuits and Crackers: Formu-
lation, Processing and Characteristics; Flatbreads of the World;
Sour Dough Technology).
sourced from fish; they are now becoming available in grains
via new genotypes of oilseeds such as canola (Figure 13) (see
articles Breeding of Grains: Canola/Rapeseed: Genetics and
Breeding; Fats: Healthy Fats and Oils; Lipid Chemistry; The
Oilseeds: Canola: Overview).

Celiac Disease: Going Gluten-Free or Not

Different Types of Fats
The various species of oilseeds provide edible oils for many of
our food sources. An example is margarine. In its processing,
some trans fats may be produced; being unnatural, trans fats
are suspect and thus discouraged. Polyunsaturated fatty acids
in our diet are considered superior to fully saturated fats, which
come mainly from animal sources. Grains are major sources of
polyunsaturated fatty acids, such as oleic acid, linoleic acid,
and linolenic acid. Some of these are essential in our diet.
Especially desirable in our diet are the long-chain omega-3
oils (eicosapentaenoic acid and docosahexaenoic acid), mainly
Only children get celiac disease and they can grow out of it
anyway. No and no!
A gluten-free diet will make you feel better. Wrong or not
necessarily.
A gluten-free diet is damaging to your health. Wrong!
A gluten-free diet will cause you to miss essential nutrients.
Maybe.
A gluten-free diet means that I cannot eat buckwheat. Wrong!
At least we know that coffee and milk are gluten-free. Not
necessarily.
 SALAD FRUIT YOGURT

Ca K Fe YOGURT

Zn Mg Mn Ca Mo K Mg
BREAD

P P I Co Na
P K Fe
FRUITS
Na CI Ca
K Mg Cu Zn
Mn
P

BUTTER

Ca Mg Na

K Co P

Se Cu

BACON AND Ca Fe K
EGGS
Na P Zn
BACON

P Cl Mg Cu

Na K S EGGS
COFFEE
BLACK PEPPER Se Fe Ca P Cr
K
Cr K S Zn Co K

EUROPEAN AND AMERICAN BREAKFAST

Figure 9 Sources of the chemical elements in breakfast. The elements carbon, oxygen, hydrogen, and nitrogen are not included because they are in all
these foods. Reproduced from Bunpei Y (2012) Wonderful Life with the Elements. San Francisco, CA: No Starch Press.

Fats, Oils & Sweets KEY

Fat (naturally occurring and added)
USE SPARINGLY
Sugars (added)
These symbols show fats and added sugars in foods.

Milk, Yogurt & Meat, Poultry, Fish, Dry Beans,

Cheese Group Eggs & Nuts Group
2-3 SERVINGS 2-3 SERVINGS

Vegetable Group Fruit Group

3-5 SERVINGS 2-4 SERVINGS

Bread, Cereal,
Rice & Pasta
Group
6-11
SERVINGS

Figure 10 Nutritional guide pyramid provided by the US Department of Agriculture.

10 THE BASICS | The Grains that Feed the World

Why the Gluten-Free Diet?
People who are diagnosed with celiac disease must adhere to a
gluten-free diet. Celiac disease is a disease affecting all ages; the
only treatment, a gluten-free diet, is a lifelong commitment.
There are other conditions, less well characterized, for which
benefit has been obtained by the omission of gluten from the
diet. In these cases, perhaps, a gluten-free diet will make you
feel better. However, beyond these cases, gluten-free diet has
become a dietary fad with many adherents. The inferior quality
of much of the gluten-free bread available is a testament to the
value of gluten in bread making. Nevertheless, there are many
substitutes for the function of gluten in the production of
gluten-free foods (Figure 14).
Strict adherence to a gluten-free is difficult given the use of
wheat, rye, triticale, and barley in many processed foods. In the
process of avoiding gluten, it may be possible that some normal
nutrient sources are also avoided with the consequent risk of
missing out on some essential nutrients. But overall, it is unlikely
that eating gluten-free diets will be damaging to your health (see
articles Appendix 3: Grains, Foods, and Ingredients Suiting

Figure 11 Nutritional guidelines as a dinner plate provided by the

Australian government. Figure 13 Canola in flower in Scotland.

sugars, fats

Acetyl-CoA

more oxaloacetate citrate more

oxidized reduced

malate [cis-aconitate]

fumarate isocitrate

succinate [oxalosuccinate]

succinyl-coA ketoglutarate

Figure 12 The Krebs cycle via which fats and sugars are broken down to CO2 releasing energy that is stored in compounds such as adenosine
triphosphate.

Figure 14 The gluten-free symbol used to indicate foods with
<20 ppm of gluten protein from wheat, rye, triticale, and barley.
Buckwheat, however, is a misnomer; it is unrelated to wheat and it is
satisfactory for people who must have a gluten-free diet (see article The
Legumes and Pseudocereals: Buckwheat: Overview).
Gluten-Free Diets for Celiac Disease; Food Grains: Intolerance,
Allergy and Diseases: Celiac Disease; Cereal Allergens; The Glu-
ten-Free Diet, the Appendix on eating gluten-free).
Even milk and coffee may not be gluten-free. Various sub-
stitutes for dairy milk involve the use of grains. The most
obvious is soy milk, which is intrinsically gluten-free as the
soya bean is unrelated to wheat. In addition, rice, oats, and
other cereals have been used to make milk substitutes, having
the dietary advantage of being lactose-free. Rice milk is gluten-
free. Oats is a satisfactory grain for most people with celiac
disease, but intake of this borderline cereal should be limited;
the equivalent of less than one cup per day is recommended
in this encyclopedias appendix (see articles Appendix 3:
Grains, Foods, and Ingredients Suiting Gluten-Free Diets for
Celiac Disease) on gluten-free diets for celiac disease.
There are various roasted-grain beverages marketed as a
caffeine-free alternatives to coffee. These coffee substitutes
may also be known as grain coffee or ersatz coffee. During
World War II, roasted wheat and barley were used as a basis
for coffee. Commercial products provide a powder form to be
reconstituted with hot water, for example, Nestle Caro.
Grains and General Nutrition
There are dietary imbalances in the cereal grains. Partly true!
Spinach is the best source of iron in our diet, because we
grew up with Popeye the Sailor Man who ate up all his
spinach. Wrong!
Dietary Imbalances?
The proteins of the major cereal grains can be criticized
for having a balance of essential amino acids that does
not fully conform to our dietary requirements. That is so.
On the other hand, the cereal grains provide an excellent
balance of proteins, carbohydrates, and fats. Some cereals
are partly deficient in some of the essential amino acids,
particularly lysine, methionine, isoleucine, and tryptophan.
THE BASICS | The Grains that Feed the World 11
In practice, these imbalances are a problem only for some
populations where there is scarce supply of complementary
food sources and also in the case of animal feeds. In con-
trast to the cereal grains, the oilseed and legume grains
more than compensate for the cereals with respect to these
essential amino acids.
Spinach?
In his comic strip, Popeyes superhuman strength was
attributed to the iron of his spinach diet, consumed by the
canful. There is a statue of Popeye in Crystal City, Texas,
commemorating his contribution to increasing the consump-
tion of spinach by 33% at about the time he was invented. In
the meat-scarce years of World War II, this was an encouraging
propaganda, but it was fraudulent. (According to Mould RF
(1984) Moulds Medical Anecdotes. Bristopl, UK: Adam Hilger
Ltd, p. 33.) The original researchers displaced the decimal
point, giving a tenfold overestimate of the iron content for
spinach. In fact, spinach has about 30 mg of iron per kilogram.
The population would have done better to concentrate on
a grain-based diet, such as wheat germ with an iron content
of about 100 mg kg1 (see articles Appendix 1: Nutrient-Com-
position Tables for Grains and for Grain-Based Products; Grain
Composition and Analysis: The Composition of Food Grains
and Grain-Based Products, the Appendix of Grain
Composition).
In Summary
Given that grains comprise such a major part of our diet, it
would be good to be able to assert that no food grains
are poisonous. However, that is not completely true, because
there is the very unusual exception of the Calabar bean; see
Figure 15. With that possible exception, it is certainly true that
the wide family of food grains is responsible directly or indi-
rectly for most of the food we eat.
Given that current world production for all grains equals
about a kilogram of grain per person per day, how can the
worlds nations work toward the elimination of malnutrition?
Exercises for Revision
How is it possible to adopt a critical attitude to assessing
nutritional information about grains presented in the
media?
What procedures should be followed to seek and research
authentic information?
Select topics, covered in this article, that are relevant to
you personally. Describe what benefit (or otherwise)
that you have obtained from information provided in
this article and from relevant articles in the Encyclopedia
of Food Grains.
Describe how grain-based foods provide the major catego-
ries of nutrients needed in our nutrition. In which cases
might certain grain products be nutritionally deficient?
Describe the categories of people who really need gluten-
free foods. Who else can be expected to benefit from gluten-
free foods?
12 THE BASICS | The Grains that Feed the World

Exercises for Readers to Explore the Topic Further

Select a few of the topics presented in this article; for these,

contrast some popular misconceptions with the authentic
concepts presented in the cross-referenced articles in the
encyclopedia and in other reliable sources, including pri-
mary scientific research articles.
Investigate some less well-known grains (e.g., chia, guar bean,
quinoa, freekeh, and macha wheat) and compare their poten-
tial benefits (in production, utilization, and nutrition) with
those of the more popular species (e.g., wheat, rice, maize, and
soy).
Investigate the routes of the major food nutrients (e.g., pro-
teins, carbohydrates, fats, and vitamins) in human digestion.
How do these compounds differ in the ways our bodies use
them?
Investigate the world production of GMO grains. What
differences are there in the regions of their production,
processing, and consumption?

Further Reading
Harlan JR (1992) Crops and Man. Madison, WI: American Society of Agronomy.
Mangelsdorf PC (1966) Genetic potentials for increasing yields of food crops and
animals. Proceedings of the National Academy of Sciences 56: 370375.
Stoskopf NC (1985) Cereal Grain Crops. Reston, VA: Reston Publishing Company.

Figure 15 The Calabar bean is poisonous because it contains

physostigmine, a reversible cholinesterase inhibitor alkaloid. It was the
likely cause of death for a Nigerian boy in London; the death has been
attributed to Nigerian witchcraft.
 Course Structures: Based on EFG Articles
H Corke, The University of Hong Kong, Hong Kong, Pokfulam, China; Hubei University of Technology, Wuhan, China
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Suggested guidelines are presented for the use of the Ency-
clopedia of Food Grains as a resource for teaching undergrad-
uate courses in food, nutrition, and agriculture.
The Encyclopedia of Food Grains is a good starting source for
authoritative material for students preparing for further in-
depth research on a topic, essays, presentations, and other
project work.
Learning Objectives
To view Encyclopedia of Food Grains as a resource for under-
graduate and graduate coursework.
To understand how Encyclopedia of Food Grains chapters can
fit additional diverse teaching and learning needs, such as
for student research project-based work and for industry-
focussed training.
Introduction
One of the special features of this new Encyclopedia of Food Grains
is the effort made to maximize its usefulness as a teaching
resource, particularly at the undergraduate level but also for
graduate-level training and industry-specific technology courses.
Education in hands-on technology-based fields such as food
science and nutrition is not all about providing suitable material
for the instructor to use in preparing lectures or for the student
to read in preparation for examinations. Modern education is
less concerned about the acquisition of specific knowledge and
more about the process of learning and inquiry, such that the
student will be prepared for lifelong learning and innovation.
The knowledge content of a subject for example, the nutri-
tional potential of a grain may change, but a well-prepared
student will always be able to seek out new information and
new ways to integrate and use the information, if they have a
sound understanding of basic concepts.
A Core Technology Course: Cereal Science/Grain
Science/Grain Processing
I teach an upper-level undergraduate course at the University of
Hong Kong called Grain Production and Utilization, which covers
Global grain production and consumption. International grain
trade. Wheat: flour milling, dough rheology, the baking process
and baking quality, quality of Asian products including steamed
bread and noodles, gluten. Rice: nutritional quality and consumer
preferences. Maize: products of wet-milling, animal feed develop-
ment. Biofuels.
Having taught this course for 20 years, I can explain in more
depth the underlying concepts and ideas. In my experience,
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00061-9
students in big city Hong Kong are surprisingly ignorant about
the basics of cereal and other grain production and usage, even
though they live facing the giant agricultural expanses of main-
land China. Many will be unclear on the basic formulation of
baked bread (guessing rice instead of wheat), and most will not
know about yeast or other leavening agents, why their favorite
rice has the texture it does, or why bioethanol is not going to
replace petroleum-based fuels.
In addition, most students will not become professional food
technologists and will hence not be interested in an engineering
unit operations approach to cereal processing. I use the theme
Food sufficiency and the global future to try to convince the
students that food grains are at the heart of human economic life,
from bread and rice to sweeteners, meat (via animal feeding), soy
proteins, and many more. The emphasis in this course is on the
interactions among technology and markets in producing,
processing, and distributing grain products.
The basic principles are interesting for such students as they
come to realize at first how little they know about the products
they eat every day rice, bread, steamed bread, noodles, etc. I tell
them that I am worried that after taking a course like this, their
relatives (usually a know-it-all uncle) may ask them something
like So you learned about grains at university, are instant noo-
dles as bad as everyone says? Everybody listen now! The expert is
going to speak. The student needs a core of factual knowledge to
be able to answer questions like that. Another major part of the
course concerns the global-scale (with an emphasis on Asia)
patterns of economic growth and land utilization that determine
the production, import, and export of food. The student should
be able to think about issues such as If oil costs $40 a barrel, or
$200 a barrel, who will eat and who will starve? Can agriculture
(bioethanol and biodiesel) provide meaningful contributions to
fuel production? Where will the grains for meat production for
China come from? These types of question are addressed on a
regular basis in the media, at least in the business context. For
students who will go on to become bankers, or personal inves-
tors, or just consumers, the ability to read such media reports
with a higher level of insight can prove very useful.
An essential part of the course is the requirement for a
hands-on lab-based group project. The theme will vary from
year to year, but a typical instruction would be
Conduct a laboratory investigation into any aspect of grain quality
measurement or the effect of ingredients on a process. The preferred
focus this year is WHEAT QUALITY BAKING or STEAMED BREAD
or NOODLES NUTRITIONAL IMPROVEMENT. For example, the
effect of protein or fiber or other additive on quality as measured by
objective means (e.g., color, texture) and by sensory evaluation.
Some attempt at OPTIMIZATION and some attempt at SENSORY
TESTING should be made. Multiple small-scale baking machines or
noodle machines can be used to design an experimental with mul-
tiple treatment comparisons.
Clearly, the students can use a resource such as Encyclopedia
of Food Grains to determine how to make their product of
13
14 THE BASICS | Course Structures: Based on EFG Articles
interest, to understand the likely effect of varying formulations,
and to seek out interesting nutritional problems and feasible
additives to make an interesting project. The use of small-scale
processing equipment has the advantage of allowing students
to make multiple samples of a product bread, steamed bread,
or noodles and with the repetition, they can remember better
the basics and develop the intuition to make intelligent adjust-
ments to the process. By observing the work of their fellow
groups in the class, and by listening to the final project pre-
sentations of all groups (in a formal half-day final symposium),
they become well acquainted with a diversity of topics.
Minimum expectations at the end of such a course would
be the ability to discuss freely, with good technical knowledge
and comparative insight, such issues as the following:
What are the differences among rice, wheat, maize, and
barley?
What climatic factors influence production of the major
cereal grains?
What are the major food uses for each?
How many fermented and distilled products are made from
grains?
To what extent are grains used for industrial products?
What limits substitution of one for the other?
What are the roles of cereal starch, oil, and protein?
What is the nutritional contribution of cereals to the
human diet?
How does meat production depend critically on grain
production?
Who will feed China?
Suitable Encyclopedia of Food Grains coverage that would support
the teaching elements of this course could include the following:
Any overview articles such as those focussing on wheat,
maize, rice, and soy.
Material on grain production and consumption at the
global and regional levels.
For selected health topics, individual topics such as Functional
Foods: Dietary Fiber, Prebiotics, Probiotics, and Synbiotics.
Elements of starch structure and functionality and analysis
of quality would help in later study of starch quality-based
products such as noodles. Protein chemistry of cereal grains
is central to any study of cereal science.
For processing wheat, an overview on wheat could be sup-
plemented with selected topics according to the instructors
and students interest, such as, on breads and on noodles.
Understanding would be strengthened by concept material
such as on sensory testing and noodle quality.
Similarly for rice, basic ideas on processing could be sup-
ported according to student need along, for example, more
in-depth genetic material or perhaps additional material on
eating quality or on the diverse uses of rice.
For maize, it is important for students to understand at least
wet-milling uses and to have a general view on animal feed.
Traditionally, soy bean has not been considered part of
grain in English language materials emphasizing cereal
grains. However, soy bean is classified with grain in Chi-
nese, and it is key agricultural product for worldwide trade,
animal feed production, and processing into food ingredi-
ents. A student must know about soy processing, and about
the use of soy concentrates and isolates.
Kansas State University has a graduate-level course Cereal Sci-
ence, with the simple course description The characteristics of
cereals, legumes, their components, and their processing to
foods. Clearly, the subject area coverage overlaps considerably
with my course, described in detail earlier, and it would be up
to the individual instructor for any such course to set out the
precise learning objectives and to select and recommend par-
ticular sections of Encyclopedia of Food Grains that would be
most helpful for the student. At graduate level, even in-depth
study of particular areas of emphasis, such as genetics and
breeding for quality, or international uses of grains, or frac-
tionation of grain, or health uses of grains, could easily be
supported by Encyclopedia of Food Grains.
Science Requirement or Reading-Intensive
Introductory Courses: Feeding the World
Most undergraduate degrees have a requirement for cross-
disciplinary knowledge, which can be developed by, for exam-
ple, requiring nonscience students to take science-based
courses (Rocks for Jocks being the famous parody of an easy
geology course designed for athletes). Another typical require-
ment is for a first year seminar-style course that imparts a
university-level way of thinking, of using sources, and of inten-
sive research and reporting. In many cases, this may be the
students first experience of small-group discussion and of
academic standards for writing and citation. In any case, such
courses should be accessible to students with little science
background yet challenging for those students who do have
strong science training. Food and nutrition provide in many
ways ideal material for such courses. The material is intrinsi-
cally interesting and accessible to students, and valuable
insights can be added from almost any perspective, from chem-
istry to economics, history, or anthropology.
I run a freshman-level course as part of the breadth require-
ment for our degree programs called Feeding the World. In a
sense, this is a world agriculture course, where students learn
about the key inputs in agriculture (e.g., energy, water,
chemicals, and knowledge) and examine the scale of world
production and its sustainability in face of constraints such as
excessive waste. The course description reads
Continuing human population increases, competition for water
supplies, and concern about energy prices have led to profound
pessimism about long-term food supplies. Already a billion people
go hungry every day. This course will offer an in-depth look at key
issues in global food sufficiency, food production, food distribution,
prospects and constraints. It is accessible to students from all fields
of study and academic backgrounds. You will develop an integrated
technical, economic and political understanding of the global food
supply crisis. You will be equipped to understand and appreciate
media reports related to this issue in your lives as informed and
influential citizens. We consider the technological, political, eco-
nomic and ethical dimensions of the global food supply enterprise.
The capacity of modern agriculture to produce food, and the capac-
ity of the political system to distribute food globally will be exam-
ined. The idea is to give an academically rigorous course that will
cross disciplinary boundaries and help develop your analytical skills
and general understanding of an important topic. The course is open
to students of all backgrounds, and hopefully this diversity can
enhance the classroom environment. The level of science required
is quite low, and none of the assessments (for example class test) will
particularly favor students with greater science background.
The week-by-week topic coverage includes Global
Production, Focus on China, The Green Revolution, Energy,
Water, Chemical Inputs, Genetic Modification, Biofuels,
Waste, and Sustainability. Clearly, these topics can be aided

by reference to selected chapters from Encyclopedia of Food
Grains. However, a major part of the course is the student
project, for which typical guidelines might be
Identify an important topic within the general theme of Who Will
Feed Africa? [other themes could be Who Will Feed China?, or Who
Can Produce Rice?, or Can Chinese Money Feed the World?]. The
topic should be one that can be investigated using statistical data
(such as production, import and export data for crops, livestock, or
chemicals; and/or data on human health and nutrition indicators).
Collect and analyze the data (preferably drawing as far as possible on
FAOStat and OECD.Stat databases). Further research the topic using
any other appropriate sources of information and prepare a written
report in Powerpoint form. Be prepared to lead a 20 min presentation
The student is then guided to pick a very narrow and
specific topic for in-depth research, for example, the potential
for Chinese investment in rice production in Mozambique. In
order to make any sense of this, they need to develop a base of
agronomic knowledge, a sense of how transferable the breed-
ing or agronomic technologies may be, and an understanding
of rice import/export balances, quality requirements, and uti-
lization in different regions. The typical Internet search may
provide a lot of fragmented and often technically superficial
information. However, Internet sources for up-to-date infor-
mation, and articles from magazines such as the Economist on
the investment climate for agriculture, seem to fit well with the
solid agricultural, production, and processing information
provided by Encyclopedia of Food Grains.
The Diversity of Courses in a Major
I have described just two typical types of undergraduate course,
rooted in knowledge of grains and grain products. We can
examine the diversity of courses spanning a typical food
science major (Table 1), albeit one with a stronger-than-
average focus on grains. Literally, all of these courses could
potentially be drawing on Encyclopedia of Food Grains for course
material, whether in agriculture, processing, chemistry, micro-
biology, analysis, engineering, or nutrition.
A similar listing of courses for a major in agriculture/agron-
omy or nutrition would also reveal substantial overlap of
course coverage with material that could be taken from Ency-
clopedia of Food Grains.
Industry Training
Typical training for industry includes 23-day short courses
on fairly general topics, such as starch quality, Asian foods,
breakfast cereals, frozen doughs, and functional foods. These
are often aimed at technical or sales staff from related areas
moving into work on a new topic for the first time. There are
also in-house training courses run by big companies to keep
their staff up-to-date with developments in, for example, grain
processing for healthy ingredients, quality testing, or new
product development. The flexibility and diversity of materials
in Encyclopedia of Food Grains provide a good starting point for
instructors building course materials for most such training.
Exercises for Revision
Choose a relevant undergraduate course and prepare a
study plan drawing on the resources of Encyclopedia of
Food Grains.
THE BASICS | Course Structures: Based on EFG Articles 15
Table 1 Core courses for the food science and industry major,
Kansas State University
FDSCI 101 Foundations in Food Science & Industry Credits: (1)
GENAG 200 Topics in Agriculture (03) College Careers (0)
FDSCI 302 Introduction to Food Science (3)
FDSCI 305 Fundamentals of Food Processing (3)
FDSCI 500 Food Science Seminar (1)
FDSCI 501 Food Chemistry (3)
FDSCI 600 Food Microbiology (2)
FDSCI 601 Food Microbiology Lab (2)
FDSCI 690 Principles of HACCP (2)
FDSCI 727 Chemical Methods of Food Analysis (2)
FDSCI 728 Physical Methods of Food Analysis (2)
GRSC 540 Engineering Applications to Grain/Food Products (3)
GRSC 541 Engineering Applications to Grain/Food Products
Laboratory (1)
Select one
ASI 318 Fundamentals of Nutrition (3)
HN 132 Basic Nutrition (3)
Select one
FDSCI 695 Quality Assurance of Food Products (3)
FDSCI 740 Research and Development of Food Products (4)
Develop the outline for a course in Food Product Develop-
ment, and suggest a health-based theme for the students to
follow in developing a team-based new product develop-
ment project. Show how the use of Encyclopedia of Food
Grains could help in providing a starting point for project
planning.
Exercises for Readers to Explore the Topic Further
Use Encyclopedia of Food Grains to develop teaching mate-
rials for your favorite course, and share the materials online
with our website!
See also: Barley, Rice and Maize Processing: Maize: Wet
Milling; Rice Processing: Beyond the Farm Gate; Carbohydrates:
Resistant Starch and Health; Starch: Analysis of Quality; Food Grains
and Well-being: Functional Foods: Dietary Fibers, Prebiotics,
Probiotics, and Synbiotics; Food-quality Testing: Noodles: Testing
for Quality; The Application of Sensory Science to the Evaluation of
Grain-Based Foods; Grains Around the World: Grain Production
and Consumption: Overview; Non-food Products from Grains:
Cereal Grains as Animal Feed; Oilseed and Legume Processing:
Soybean: Processing; Soybean: Soy Concentrates and Isolates;
Proteins: The Protein Chemistry of Cereal Grains; The Cereal
Grains: Maize: Overview; Wheat: An Overview of the Grain That
Provides Our Daily Bread; The Oilseeds: Soybean: Overview;
Wheat-Based Foods: Breads; Cereals: Overview of Uses: Accent on
Wheat Grain; Noodles: Asian Wheat Flour Noodles.
Relevant Websites
http://www.grains.k-state.edu/undergraduate-programs/ Website of the Department of
Grain Science and Industry of Kansas State University, with listing and descriptions
of all undergraduate courses.
 The Grain Crops: An Overview
RA Graybosch, USDA-ARS, University of Nebraska Lincoln, Lincoln, NE, USA
Published by Elsevier Ltd.
Topic Highlights
Definition of a grain
Botanical classification of grain crops
Utilization of grain crops
Definition and classification of a pseudocereal
Systematic listing of grain crop species and their botanical
families
Classification of primitive grain crops
Learning Objectives
To understand the definition of grain
To appreciate the evolutionary diversity of grains and struc-
tures utilized as grains
To comprehend the difference between common and sci-
entific names and understand the need for precision and
the need for scientific names in technical literature
To have knowledge of the difference between cereals and
pseudocereals
To appreciate the diversity of grain crops and their
utilization
Introduction
Plants considered grain crops are those producing small, hard,
dry seed or fruit consumed by man or his domesticated animals
as a foodstuff or processed for food or industrial purposes.
Grain crops, as a grouping, is, however, largely artificial.
Plants producing useful grains have evolved in a number of
plant families, and these families are not always closely related.
Botanically speaking, grains themselves are heterogeneous. The
term was derived from the Latin granum meaning seed. Cereal
grains, however, actually are formed of an entire fruit (caryop-
sis), while those of other grain crops are the dry seeds of various
types of fruits including legumes (pulses), achenes (sunflowers
and buckwheats), siliques (canola), and capsules (cotton).
Grains, therefore, are the result of convergent evolution or the
development of similar structures (grains) in diverse organ-
isms. The actual genes involved in the formation of these
structures might be different in each family of grain crops.
The Grain Crops
Grain crops of significance are listed in Table 1 according to
their common English names and the equivalent botanical
names (as genus and species). When such lists are formulated,
it is valuable to arrange species in a systematic or taxonomic
fashion. This has been accomplished in Table 1 by grouping
16 Encyclopedia of Food Grains, Second Edition
included species in orders, families, and, for the two most
important families, the Poaceae and the Fabaceae, into tribes.
Listed crops may have relatives cultivated locally (see ars-grin.
gov http://www.ars-grin.gov for additional information).
Common names are the local, regional, or national names
for plants. With the multitude of spoken languages in exis-
tence, common names for many crop species often are numer-
ous. In scientific literature, common names for plants should
be used only when the scientific name is presented at first
mention of a plant species. The use of common rather than
scientific names can lead to needless confusion. Some plant
species are known, even within a single language, by a multi-
tude of common names. For example, the grain crop proso
millet (Panicum miliaceum L.) is also known by the English
common names common millet, French millet, hog millet,
broomcorn, and broomcorn millet. Also, some common
names have been applied to totally different species. In the
Americas, corn refers to Zea mays L., while in Europe, the term
has been applied to barley (Hordeum vulgare L.) or to spelt
wheat (Triticum aestivum L. subsp. spelta (L.) Thell.). The use
of the scientific name would avoid the obvious confusion that
might result from use of common names.
Due to extensive morphological variation, numerous sub-
specific names have been assigned to some grain crops and
their relatives. Subspecies names are useful as a means of
differentiating various forms of cultivated crops, especially
when distinct market classes exist for each type. For example,
common or bread wheat is classified as T. aestivum subsp.
aestivum. Two morphologically distinct forms of T. aestivum
L., club wheat (T. aestivum L. subsp. compactum (Host) Mackey)
and spelt (T. aestivum L. subsp. spelta (L.) Thell.) are grown and
marketed for different end uses. The use of the subspecies
names, in such cases, is useful as it differentiates the various
cultivated forms. Subspecies names also are useful to differen-
tiate cultivated forms of plant species from wild ones. For
example, durum wheat (T. turgidum L. subsp. durum) has sev-
eral wild, semiwild, and formerly cultivated relatives including
T. turgidum subsp. dicoccoides and T. turgidum subsp. dicoccon.
In both examples, the use of the subspecies names serves to
differentiate the morphologically different forms while retain-
ing an indication of the close genetic relationships between the
various types. Such information is useful as an indication of
the existence of close relatives of grain crops for use as gene
donors in crop improvement programs. It also is useful as an
indicator to people with dietary restrictions (e.g., those
afflicted with celiac disease) of the true genetic relationship
between grain crops.
Plants classified as grain crops are found in both classes of
the division Anthophyta (Table 1) and include representatives
of 10 orders and 11 families. The families with the largest
number of members are the Poaceae (Gramineae), also
known as the grass or cereal family, and the Fabaceae (Legu-
minosae), the legume or bean family. These two families
http://dx.doi.org/10.1016/B978-0-12-394437-5.00001-2

Table 1 The grain crops, their family groupings, and botanical names (genus and species), plus common names
Taxonomic grouping
Class: Liliopsida
Order: Cyperales
Family: Poaceae (Gramineae)
Subfamily: Pooideae
Tribe Poeae
Tribe Triticeae
Tribe Stipeae
Subfamily: Panicoideae
Tribe Andropogoneae
Tribe Paniceae
Subfamily: Chloridoideae
Tribe Chlorideae
Tribe Eragrosteae
Subfamily: Bambusoideae
Tribe Oryzeae
Class: Magnoliopsida
Order: Fabales
Family: Fabaceae (Leguminosae)
Tribe Dalbergieae
Tribe Cicereae
THE BASICS | The Grain Crops: An Overview 17
Genus Species (accepted namea) English common names
Avena Avena abyssinica Hochst. in Schimper Abyssinian oat
Avena brevis Roth Short oat
Avena nuda L. Naked oat
Avena sativa L. Oats, common oat
Avena strigosa Schreb. Black oat, small oat
Phalaris Phalaris canariensis L. Canary grass
Hordeum Hordeum vulgare L. Barley
Secale Secale cereale L. Rye
Secale
derzhavinii Tzvelev Perennial rye
X Triticosecale
Triticosecale Wittm. Triticale
Triticum Triticum aestivum L. subsp. aestivum Common wheat, bread wheat
Triticum aestivum L. subsp. compactum (Host) Club wheat
Mackey
Triticum aestivum L. subsp. spelta (L.) Thell. Spelt, farro
Triticum monococcum L. subsp. monococcum Small spelt, einkorn, farro
Triticum timopheevii (Zhuk.) Zhuk. subsp. Zanduri wheat, Georgian wheat
timopheevii
Triticum turgidum L. subsp. durum (Desf.) Husn. Durum, durum wheat
Triticum turgidum L. subsp. carthlicum (Nevski) Persian wheat
A. & D. Love
Triticum turgidum L. subsp. dicoccon (Schrank) Emmer
Thell.
Triticum turgidum L. subsp. polonicum (L.) Thell. Polish wheat
Triticum turgidum L. subsp. turanicum (Jakubc.) Khorasan wheat
A. & D. Love
Triticum turgidum L. subsp. turgidum Cone wheat, pollard wheat, rivet wheat
Achnatherum Achnatherum hymenoides (Roem. & Schult.) Indian ricegrass
Barkworth
Coix Coix lacryma-jobi L. Coix, Jobs tears
Zea Zea mays L. Maize, Indian corn, corn
Sorghum Sorghum bicolor (L.) Moench Sorghum
Pennisetum Pennisetum glaucum (L.) R. Br. Pearl millet
Panicum Panicum miliaceum L. Proso millet, common millet, French millet,
hog millet, broomcorn millet
Panicum hiritcaule J. Presl Sowi millet, sauwi
Panicum sumatrense Roth Little millet, blue panic
Setaria Setaria italica (L.) P. Beauv. Foxtail millet, Hungarian millet, Italian millet
Echinochloa Echinochloa esculenta (A. Braun) H. Scholz Japanese millet, Japanese barnyard millet
Echinochloa frumentacea Link Sawa millet, sawa, billion dollar grass
Paspalum Paspalum scrobiculatum L. Kodo millet
Digitaria Digitaria exilis (Kipp.) Stapf Fonio, white fonio, white fonio millet
Digitaria iburua Stapf Black fonio
Urochloa Urochloa deflexa (Schumach.) H. Scholz Guinea millet, animal fonio
Eleusine Eleusine coracana (L.) Gaertn. Finger millet, birdsfoot
Eragrostis Eragrostis tef (Zuccagni) Trotter Teff
Oryza Oryza sativa L. Rice
Oryza glaberrima Steud. African rice
Zizania Zizania palustris L. Northern wild rice
Zizania aquatica L. Wild rice, zizania
Arachis Arachis hypogaea L. subsp. hypogaea Virginia peanut
Arachis hypogaea L. subsp. fastigiata Waldr. Peanut, Spanish or Valencia types
Cicer Cicer arietinum L. Chickpea
(Continued)
18 THE BASICS | The Grain Crops: An Overview

Table 1 (Continued)

Taxonomic grouping Genus Species (accepted namea) English common names

Tribe Genisteae Lupinus Lupinus luteus L. Yellow lupin

Lupinus angustifolius L. Blue lupin
Lupinus albus L. White lupin
Tribe Indigofereae Cyamopsis Cyamopsis tetragonoloba (L.) Taub. (L.) Taub. in Guar
Engl. & Prantl
Tribe Phaseoleae Cajanus Cajanus cajan (L.) Huth Pigeon pea
Canavalia Canavalia ensiformis (L.) DC. Jack bean, horse bean
Canavalia gladiata (Jacq.) DC. Sword bean
Glycine Glycine max (L.) Merr. Soybean, soya
Lablab Lablab purpureus (L.) Sweet Hyacinth bean
Macrotyloma Macrotyloma geocarpum (Harms) Marechal & Kerstings groundnut
Baudet
Mucuna Mucuna pruriens (L.) DC. Velvet bean
Phaseolus Phaseolus acutifolius A. Gray Tepary bean
Phaseolus coccineus L. Runner bean, scarlet runner bean
Phaseolus lunatus L. Butter bean, lima bean
Phaseolus vulgaris L. Dry edible beans, kidney bean, navy bean,
wax bean, green bean, field bean, etc.
Psophocarpus Psophocarpus tetragonolobus (Stickm.) DC. Winged bean
Vigna Vigna aconitifolia (Jacq.) Marechal Moth bean
Vigna angularis (Willd.) Ohwi & Ohashi Adzuki bean
Vigna radiata (L.) R. Wilczek Mung bean
Vigna mungo (L.) Hepper Black gram
Vigna subterranea (L.) Verdc. Bambara groundnut
Vigna umbellata (Thumb.) Ohwi & Ohashi Rice bean
Vigna unguiculata (L.) Walp. Black-eyed pea, cowpea
Tribe Fabeae Lathyrus Lathyrus sativus L. Chickling pea, chickling vetch
Lens Lens culinaris Medik. Lentil
Pisum Pisum abyssinicum A. Braun Abyssinian pea
Pisum sativum L. subsp. sativum Pea, field pea, sugar pea

Pisum sativum L. subsp. asiaticum Govor. Asiatic pea

Order: Caryophyllales
Family: Amaranthaceae
Family: Chenopodiaceae
Family: Chenopodiaceae
Order: Asterales
Family: Compositae
(Asteraceae)
Order: Lamiales
Order: Euphorbiales
Euphorbiaceae
Order: Linales
Family: Linaceae
Order: Scrophulariales
Family: Pedaliaceae
Order: Polygonales
Family: Polygonaceae
Order: Capparales
Family: Brassicaceae
(Cruciferae)
Order: Malvales
Family: Malvaceae
a
From USDA-ARS grin taxonomy for plants (http://www.ars-grin.gov/cgi-bin/npgs/html/index.pl?languageen).
Pisum sativum L. subsp. transcaucasicum Govor. Caucasus Mountain pea
Vicia Vicia ervilia (L.) Willd. Bitter vetch
Vicia faba L. Broad bean; fava, fava bean
Amaranthus Amaranthus hypochondriacus L. Grain amaranth; princes feather
Amaranthus caudatus L. Amaranth, Inca wheat
Amaranthus cruentus L. Purple amaranth, red amaranth, amaranth
Chenopodium Chenopodium quinoa Willd. Quinoa
Chenopodium pallidicaule Aellen Canihua
Carthamus Carthamus tinctorius L. Safflower
Guizotia Guizotia abyssinica (L.f.) Cass. Niger thistle, niger
Helianthus Helianthus annuus L. Sunflower
Salvia Salvia potus Epling Chia
Ricinus Ricinus communis L. Castor bean
Linum Linum usitatissimum L. Linseed, flax, linola
Sesamum Sesamum indicum L. Sesame
Fagopyrum Fagopyrum esculentum Moench Buckwheat, Japanese buckwheat
Fagopyrum tataricum (L.) Gaertn. Tartary buckwheat
Brassica Brassica napus L. Canola, rape, oilseed rape
Brassica nigra (L.) Koch Black mustard
Brassica juncea (L.) Czern Brown mustard, sarson
Crambe hispanica L. subsp. abyssinica (Hochst. ex Crambe
R. E. Fr.) Prina
Gossypium Gossypium arboreum L. Tree cotton
Gossypium barbadense L. American Pima cotton, Sea Island cotton
Gossypium herbaceum L. Arabian, Levant or Maltese cotton
Gossypium hirsutum L. American cotton, upland cotton

contain over about 70% of the species considered grain crops.
Five species alone, including four grasses, wheat (T. aestivum
and T. turgidum subsp. durum), maize (Z. mays L.), rice (Oryza
sativa L.), and barley (H. vulgare L.), and the legume soybean
(Glycine max (L.) Merr.) also provide over 70% of the worlds
metric tonnage of food from annual plants.
The Grasses
The grass family (Poaceae) contains the largest and, in terms of
annual production, most important grain crops, the cereals.
The Poaceae is one of the largest plant families, commonly
divided into subfamilies, each, in turn, containing one or more
tribes. Cereal grains (Table 1) are found in the subfamilies
Pooideae, Bambusoideae, Chloridoideae, and Panicoideae,
with the Pooideae and Panicoideae containing the largest
numbers of grain crop species. Early systems of grass taxonomy
relied heavily on morphological features, especially spikelet
and inflorescence morphology. However, beginning in the
1950s, information regarding chromosome numbers, embryo
structure, leaf anatomy, interspecific hybridizations, and types
of photosynthetic systems began to accumulate, and it became
clear that the early arrangement of grass genera into subfam-
ilies and tribes was largely artificial. In the 1960s, grass taxon-
omy was completely revised. Many earlier volumes still in wide
use (e.g., Hitchcock and Chases Manual of the Grasses of the
United States) reflect the original groupings of grass genera.
More recent references (e.g., Watson and Dallwitzs Grass
Genera of the World http://delta-intkey.com/grass/), followed
herein, should be consulted for proper tribal affiliation of grass
genera.
Wheat and other members of the tribe Triticeae (subfamily
Pooideae) are among the most important grain crops. Com-
mon or bread wheat, T. aestivum L. subsp. aestivum, and durum
wheat, T. turgidum (L.) subsp. durum (Desf.) Husn., are the two
most widely cultivated. Two additional subspecies of T. aesti-
vum, T. aestivum L. subsp. compactum (Host) Mackey (club
wheat) and T. aestivum L. subsp. spelta (L.) Thell. (spelt or
farro), also are cultivated. Club wheats are used to produce
pastry and other low protein flours, and spelt is grown largely
as an heirloom crop, often in organic production systems.
Some authors consider these to be separate species, but they
both differ from typical bread wheat only in some aspects of
spike morphology controlled by one or a few genes, and the
forms will all freely hybridize, so recognition as separate spe-
cies is unwarranted. T. monococcum L. subsp. monococcum
(small spelt, einkorn, or farro) was once grown as a grain
crop in most states bordering the Mediterranean Sea and
throughout Europe. T. timopheevii (Zhuk.) Zhuk. (Georgian
or zanduri wheat) was cultivated in what is now the nation
of Georgia. Both T. monococcum and T. timopheevii have all but
disappeared from modern cultivation. Several additional sub-
species of T. turgidum have been cultivated in historical times,
but either their importance has diminished, or they are culti-
vated in local, isolated areas as relic crops. These include T.
turgidum L. subsp. carthlicum (Nevski) A. & D. Love (Persian
wheat), T. turgidum L. subsp. dicoccon (Schrank) Thell.
(emmer), T. turgidum L. subsp. polonicum (L.) Thell. (Polish
wheat), T. turgidum L. subsp. turanicum (Jakubc.) A. & D. Love
THE BASICS | The Grain Crops: An Overview 19
(Khorasan wheat), and T. turgidum L. subsp. turgidum (cone,
pollard, or rivet wheat). Other important members of the tribe
Triticeae include barley (H. vulgare L.) and rye (Secale cereale L.).
Triticale (X Triticosecale Wittm.) is a man-made crop, grown
primarily not only as forage but also as a grain in some parts of
the world. Triticales were developed via hybridizations between
common or durum wheats and rye. Octoploid triticales contain
all the genes of hexaploid wheat and rye, while hexaploid triti-
cales contain the genes of tetraploid durum wheat and rye.
Secale
derzhavinii Tzvelev, another man-made crop, has been
developed as a perennial grain crop. Its cultivation is limited.
The oats, Avena, are more distantly related, being placed
along with the wheats in the subfamily Pooideae, but in a
different tribe, the Poeae. Common oat, Avena sativa L., is the
most commonly cultivated species. Its importance has dimin-
ished in historical times with the change from animal-driven to
mechanized agriculture, but oats are still an important grain
crop for human consumption, especially in the northern hemi-
sphere. Other species of Avena have been grown as grain crops,
but they either are no longer cultivated or have been reduced to
the status of relic crops. These included A. abyssinica Hochst. in
Schimper (Abyssinian oat), A. brevis Roth (short oat), A. nuda
L. (naked oat), and A. strigosa Schreb. (black or small oat).
Phalaris canariensis L., another member of the tribe Poeae,
primarily is grown as a source of feed for caged birds.
Maize, sorghum, and Jobs tears
Maize (Z. mays L.) and sorghum (S. bicolor (L.) Moench.) are
both members of the subfamily Andropogoneae. Jobs tears
(Coix lacryma-jobi L.), rarely cultivated as a grain, but more
frequent now as an ornamental crop, also is a member of this
tribe. Many morphological types of both maize and sorghum
exist; in both cases, however, only a single species and subspe-
cies are recognized.
The millets and teff
The millets are a taxonomically confusing group. The term
millet evidently has been applied to any grass with a small
round seed. Thus, the term is used to describe a number of, at
times, only distantly related species. Most of the millets are
placed in the subfamily Panicoideae, tribe Paniceae. The most
important millet in terms of annual production is pearl millet
(Pennisetum glaucum (L.) R.Br.), most commonly grown in sub-
Saharan Africa and in India. The remaining millets often are
termed minor millets. Of these, proso millet (P. miliaceum L.)
is the most widespread, grown as a grain crop throughout Asia
and into Eastern Europe. In North America, it is largely grown
as a feed for caged and domestic birds. Several additional
millets are grown, primarily in Asia. The following also are
members of the tribe Paniceae: Setaria italica (L.) P. Beauv.
(foxtail millet), Echinochloa esculenta (A. Braun) H. Scholz
(Japanese millet), E. frumentacea Link. (sawa millet), Paspalum
scrobiculatum L. (kodo millet), Digitaria exilis (Kipp.) Stapf
(fonio or white fonio), D. iburua Stapf (black fonio), and
Urochloa deflexa (Schumach.) H. Scholz (Guinea millet). Eleu-
sine coracana (L.) Gaertn, known as finger millet or birdsfoot,
actually is a member of the subfamily Chloridoideae, tribe
Chlorideae, indicating that it is more closely related to teff
than it is to the rest of the millets. Teff (Eragrostis tef (Zuccagni)
Trotter), subfamily Chloridoideae, tribe Eragrosteae, is
20 THE BASICS | The Grain Crops: An Overview
cultivated as a grain crop in Ethiopia and as a forage grass
elsewhere. It is the only cultivated grain species from a fairly
large genus of grasses.
Rice and its relatives
Rice (O. sativa L., tribe Oryzeae) is the major food of nearly
50% of the worlds population. Two forms or races of rice are
recognized, indica types (race indica), grown in the tropics,
and japonica types (races japonica and javonica), the temperate
forms. These types are not recognized as subspecies. African
rice (O. glaberrima Steud.) long has been cultivated in Western
and Central Africa, but it has largely been replaced by O. sativa.
Zizania palustris L., northern wild rice, and Z. aquatica L. also
members of the tribe Oryzeae, are cultivated and gathered from
wild populations in North America. Oryza and Zizania are the
only grain crops assigned to the subfamily Bambusoideae.
Indian ricegrass (Achnatherum hymenoides (Roem. & Schult.)
Barkworth), formerly was gathered as a grain crop by Native
Americans, and there have been some recent attempts to culti-
vate and market it. Indian ricegrass is so named for its super-
ficial morphological resemblance to rice, but it actually is a
member of the subfamily Pooideae.
The Legumes
The Fabaceae (legume or bean family) is second only to the
Poaceae in terms of economic importance to man. This large
family is divided into three subfamilies, the Papilionoideae,
Caesalpinioideae, and Mimosoideae. Some authors treat these
as separate families. All of the grain legumes are members of
the Papilionoideae. Most grain legumes are members of the
tribes Phaseoleae and Vicieae, although some also are assigned
to the tribes Dalbergieae, Cicereae, Genisteae, and Indigofereae
(Table 1).
Soybean, or soya (Glycine max L.), is the most important, in
terms of annual metric tonnage of production. G. soja Siebold
& Zucc., the wild progenitor of soybean, is a close relative, and
the two are, at times, treated as one species. Intermediate types
exist and formerly were designated G. gracilis Skvortzov, but
they now are recognized as representatives of the subspecies G.
max subsp. gracilis (Skvortzov) Enken.
Edible dry beans are assigned to the species Phaseolus vul-
garis L. A large number of morphological variants exist for this
species, some being consumed as pulses, others for the edible
pods. Common names include dry edible beans, kidney bean,
navy bean, wax bean, green bean, and field bean. All are
considered members of the same species. Other species of
Phaseolus are cultivated, including the tepary bean, P. acutifolius
A. Gray, P. coccineus L. (scarlet runner bean), and P. lunatus L.
(butter bean or lima bean).
Arachis hypogaea L. (peanut) is widely cultivated in both the
Old and New Worlds. Two subspecies are grown, A. hypogaea L.
subsp. hypogaea (Virginia peanut) and A. hypogaea L. subsp.
fastigiata Waldr. (Spanish or Valencia peanut). Other impor-
tant grain legumes include Vicia faba L. (broad bean or fava),
Cajanus cajan (L.) Millsp. (pigeon pea), Cicer arietinum L.
(chickpea), Lens culinaris Medik. (lentil), and Pisum sativum L.
(field or sweet pea). Several subspecies of P. sativum are culti-
vated, with P. sativum L. subsp. sativum, the common garden
pea, being the predominant form. Other less common forms
include P. sativum L. subsp. asiaticum Govor. (Asiatic pea),
cultivated in Asia, and P. sativum L. subsp. transcaucasicum
Govor., cultivated in the Caucasus mountain region. Legumes
of lesser importance include Cyamopsis tetragonoloba (L.) Taub.
(L.) Taub. in Engl. & Prantl (guar), cultivated at times as a grain
but now more important as a source of guar gum. Other less
common grain legumes are three species of the genus Lupinus,
L. luteus L. (yellow lupin), L. angustifolius (blue lupin), and L.
albus (white lupin). The lupins also are grown as ornamental
plants. Widely cultivated, though of only local importance as
grain crops, are two species of the genus Canavalia (jack bean
and sword bean), Lablab purpureus (L.) Sweet (hyacinth bean),
Lathyrus sativus L. (chickling pea), Macrotyloma geocarpum
(Harms) Marechal & Baudet (Kerstings groundnut), Psophocar-
pus tetragonolobus (Stickm.) DC. (winged bean), Vicia ervilia
(L.) Willd. (bitter vetch), Mucuna pruriens (L.) DC. (velvet
bean), and several species of the genus Vigna, V. angularis
(Willd.) Ohwi & Ohashi (adzuki bean), V. aconitifolia (Jacq.)
Marechal (moth bean), V. radiata (L.) R. Wilczek (mung bean),
V. mungo (L.) Hepper (black gram), V. subterranea (L.) Verdc.
(Bambara groundnut), V. umbellata (Thunb.) Ohwi & Ohashi
(rice bean), and V. unguiculata (L.) Walp. (black-eyed pea or
cowpea).
Oilseed Crops and Pseudocereals
Several crops of various families produce grains from which
edible or industrial oils are processed. The mustard family
(Brassicaceae or Cruciferae) contains several oilseed species,
most important of which is Brassica napus L. (rapeseed or
canola). B. nigra (L.) Koch in Rohling (black mustard) and B.
juncea (L.) Czern. (brown mustard) occasionally are cultivated
for oil and to produce mustard. Crambe hispanica L. subsp.
abyssinica (Hochst. ex R. E. Fr.) Prina (crambe) is another
member of the mustard family used as a source of oil for
various industrial purposes. One member of the family Peda-
liaceae, Sesamum indicum L. (sesame), is cultivated as both a
spice and a source of sesame oil. Ricinus communis L. (castor
bean), of the family Euphorbiaceae, is the source of castor oil,
used in industrial applications. It is also infamous as the source
of the toxin ricin, a seed protein. Linseed oil is derived from
Linum usitatissimum L., a member of the Linaceae. L. usitatissi-
mum also is the source of flax. Chia (Salvia potus Epling) a
member of the mint family (Lamiaceae) is used as a source of
edible oil and consumed directly in various ways including a
topping for yogurt and as an ingredient for breakfast cereals.
Chia seed also is used to produce chia pets.
The Compositae (Asteraceae), or sunflower family, proba-
bly is the worlds largest family, in terms of number of species.
However, it contains only three members cultivated as grain
crops, Helianthus annuus L. (sunflower), Guizotia abyssinica (L.
f.) Cass. (Niger thistle), and Carthamus tinctorius L. (safflower).
Sunflower and safflower primarily are grown as oilseed crops,
but sunflower also is used as a grain for human and avian
consumers. Niger thistle also is grown for birdseed.
Cotton species (family Malvaceae, genus Gossypium) inde-
pendently were domesticated in both the Old and New
Worlds. G. herbaceum L. (Arabian or Egyptian cotton) was of
Old World origin, while G. hirsutum L. (American upland
cotton) and G. barbadense L. (American Pima cotton) and

G. arboreum L. (tree cotton) were domesticated in the New
World. Over 90% of the worlds cotton production now is
from G. hirsutum L. While primarily grown as a fiber crop,
both oil and protein are processed from cotton seeds.
Pseudocereals are nonmembers of the grass family often
milled to flour and used to produce flat breads and other
products similar to those derived from true cereals. This is
another artificial grouping, containing members of several
plant families. Two members of the Chenopodiaceae are culti-
vated, Chenopodium quinoa Willd. (quinoa) and C. pallidicaule
Aellen (canihua). The grain amaranths, Amaranthus hypochon-
driacus L., A. caudatus L., and A. cruentus L., are assigned to the
Amaranthaceae. These five species all were components of the
ancestral cultivated flora of Mesoamerica and South America.
Additional pseudocereals include Fagopyrum esculentum
Moench. (buckwheat) and F. tataricum (L.) Gaertn. (Tartary
buckwheat), both members of the Polygonaceae. Buckwheats
probably were first cultivated in Western China.
Exercises for Revision
What is the difference between a cereal and a pseudocereal?
Explain the difference between a scientific and a common
name? When should scientific names be used? What is the
point of using names that are difficult to spell and
remember?
What is the difference between a species and a subspecies?
What is a millet? Are all millets derived from the same
species?
Which are more closely related wheat and oats or wheat
and rice? Why?
Why do you suppose the majority of our grain crops derive
from only two plant families?
Exercises for Readers to Explore the Topic Further
Has the worlds biodiversity of grain crops been fully
explored? Are there under- or nonutilized species that
could be produced and consumed for nutritional,
environmental, or cultural enrichment?
Can pseudocereals be more effectively used in modern food
manufacturing? Will their increased use slow or diminish
the rate of incidence of celiac disease?
Can wild or semiwild relatives of grain crops be more
effectively exploited? Can useful genes in these species be
accessed without genetic engineering?
Could ancient crops no longer cultivated be resurrected,
and would more widespread culture and utilization
improve nutritional composition of modern diets and
lead to less environmentally damaging agricultural
systems?
See also: Agronomy of Grain Growing: Sorghum: Production and
Improvement Practices; Breeding of Grains: Barley: Genetics and
Breeding; Canola/Rapeseed: Genetics and Breeding; Lentil: Breeding;
THE BASICS | The Grain Crops: An Overview 21
Soybean: Germplasm, Breeding, and Genetics; Wheat Breeding:
Exploiting and Fixing Genetic Variation by Selection and Evaluation;
Genetics of Grains: Maize: Genetics; The Basics: Taxonomic
Classification of Grain Species; The Cereal Grains: An Overview of
the Family of Cereal Grains Prominent in World Agriculture; Cereals:
Domestication of the Cereal Grains; Rice: Overview; The Legumes
and Pseudocereals: Amaranth: Overview; Beans: Overview;
Buckwheat: Overview; Chickpea: Overview; Grain Legumes and Their
Dietary Impact: Overview; Lupin: Overview; Pea: Overview;
Pseudocereals: Overview; The Oilseeds: Oilseeds: Overview.
Further Reading
Belton PS and Taylor JRN (2002) Pseudocereals and Less Common Cereals: Grain
Properties and Utilization Potential. Berlin: Springer.
Brummitt RK and Powell CE (eds.) (1992) Authors of Plant Names. Kew, UK: Royal
Botanical Gardens.
Cronquist A (1981) An Integrated System of Classification of Flowering Plants.
New York: Columbia University Press.
Cronquist A (1988) Evolution and Classification of Flowering Plants. New York:
New York Botanical Gardens.
Duke JA (1981) Handbook of Legumes of World Economic Importance. New York:
Plenum Press.
Gould FW (1968) Grass Systematics. New York: McGraw Hill.
Hanelt P (ed.) (2001) Mansfelds Encyclopedia of Agricultural and Horticultural Crops.
Heidelberg: Springer.
Harlan JR (1992) Crops and Man, 2nd edn. Madison, WI: American Society of
Agronomy.
Hitchcock AS (1971) Manual of the Grasses of the United States, 2nd edn. New York:
Dover, revised by A. Chase.
Lawrence GHM (1951) Taxonomy of Vascular Plants. New York: Macmillan.
Mabberley DJ (1997) The Plant-Book. Cambridge: Cambridge University Press.
Pohlhill RM and Raven PH (1981) Advances in Legume Systematics. Kew, UK: Royal
Botanic Gardens.
Smartt J (1990) Grain Legumes. Evolution and Genetic Resources. Cambridge:
Cambridge University Press.
Vavilov NI (1960) World Resources of Cereals, Leguminous Seed Crops and Flax and
Their Utilization in Plant Breeding. Jerusalem: Israel Program for Scientific
Translations.
Wiersema JH and Blanca L (1999) World Economic Plants: A Standard. Boca Raton, FL:
CRC Press.
Relevant Websites
http://www.ars-grin.gov On-line searchable version of World Economic Plants:
A Standard Reference. This version includes all plant species listed in the reference
and maintained in the United States Department Agricultures National Plant
Germplasm Collection. Searches can be performed on scientific names and plant
uses or other attributes of economic importance. Also includes complete lists of
common names and synonyms.
The grass genera of the world L. Watson and M. J. Dallwitz, http://delta-intkey.com/
grass/ This website provides a complete taxonomic description of the worlds
grass genera, including information on assignment to tribes, descriptions.
http://www.iapt-taxon.org/nomen/main.php International Code of Botanical
Nomenclature accepted rules for naming plant species.
http://www.ildis.org/LegumeWeb k LegumeWeb from the International Legume Data
and Information Service World Database of Legumes, a searchable database
providing complete information on legume taxonomy, including accepted names,
assignment to tribes, references, etc.
mansfeld.ipk-gatersleben.de http://mansfeld.ipk-gatersleben.de/pls/htmldb_pgrc/f?
p 185:3:0 On-line version of Mansfelds Encyclopedia of Agricultural and
Horticultural Crops. Searchable, includes scientific names, synonyms, common
names, discussion, and references for what seems to include every plant species
ever cultivated by man.
 The Grain Chain: The Route from Genes to Grain-Based Products
CW Wrigley, University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The value-addition chain for grain starts with the breeder,
who builds suitable genes into a specific variety so that it
will provide a good yield of quality grain.
Farm management is designed to optimize yield and grain
quality, although some aspects of the growth environment
may be beyond the growers control.
After harvest, grain may be tested and segregated according
to its suitability for processing.
The next stage of processing covers a wide range of possi-
bilities, given the many grain species and the great variety of
possible processes. Some examples are given.
The emphasis is on the consumer as the ultimate person to
determine product quality by deciding what products to
buy.
Thus, there is feedback up the grain chain according to the
quality of the product or intermediate provided by the
previous person in the chain.
Learning Objectives
To achieve an understanding of all the stages leading to the
eventual consumer product, via breeding, grain production,
processing, and marketing
To gain an appreciation of the need for those involved at
each stage to satisfy the requirements of those further down
the chain, most importantly the consumer
Introduction
When you bite into a delicious pastry, do you consider where
its ingredients came from? The main component is wheat
flour, which came through the sequence of stages shown in
Figure 1. This chain of events relates to the development,
production, and processing of all grain species through their
utilization. The sequence has been called field to fork and
paddock to plate for grain-based foods, but it applies similarly
for industrial products such as adhesives, starch for paper
manufacture, animal feeds, and bioethanol as a vehicle fuel.
In some cases, the processing ends at the stage of the whole
grain, such as rice, couscous, and sweet corn.
This grain chain is a value-adding sequence. In the case of
the pastry, which may have cost you about US$3, the major
component (flour) came from the processing of about US
$0.03 worth of grain (based on the per tonne price of wheat).
The value addition from the farm gate to the pastry shop or
patisserie was a 100-fold. In the case of a pastry bought in some
countries, this sequence may occur over long distances involv-
ing rail and sea transport from farm production to the sites of
processing and retail sale.
22 Encyclopedia of Food Grains, Second Edition
On the other hand, the whole sequence may occur in one
village for a family on subsistence farming, with some grains
being cooked and consumed as is by the family and other
grains being processed in a village-owned mill. In place of the
marketing stage of the chain in Figure 1, there may be the
interchange of various types of food or labor. Nevertheless,
even in this case, there will be the aspect of consumer reaction
affecting decisions about what species and type of grain to sow
in the subsequent season.
A circle format (Figure 2) is an alternative representation of
the value-addition sequence because it emphasizes the feed-
back function, primarily the communication from the con-
sumer back to the breeder, who must produce varieties with
quality traits that suit the specific consumer products. But in
addition, many other feedback functions are needed:
From the grower to the breeder (I need disease-resistant,
high-yielding varieties)
From the rice-mill operator to the grower (I need rice grain
that mills well with a minimum of broken grains)
From the maltster to the grower (I need barley that germi-
nates uniformly)
From the hummus maker to the grower or grain trader
(I need good chickpea grain for making my hummus)
From the miller to the grower (I need clean grain that gives
a high flour yield)
From the baker to the miller (I need flour that forms a
strong extensible dough that makes good bread)
From the retailer to the food processer (I need products
that look attractive, that taste good, and that sell well)
The Grain-Supply Sequence Reflected in This
Encyclopedia
The arrangement of articles in this four-volume encyclopedia
reflects the grain-supply chain, but in reverse. Volume 4 covers
grain production and genetics. Volume 3 describes grain-based
products and their processing methods. The consumer and
nutritional topics are featured prominently in Volume 2,
these being issues of special public interest. Volume 1 provides
a general overview of grains under the title The World of
Grains. Issues of grain processing and products were consid-
ered to warrant a full volume (3). Volume 4 features most of
the reverse grain chain of marketing and grading, transport,
agronomy, breeding, and genetics.
Breed: The Production of Improved Varieties Is a Critical
Contribution of Grain Science
Plant breeding has developed during the past century as the
means of producing suitable genotypes, selecting them on the
basis of the required attributes, after the use of crossbreeding to
expand the genetic diversity available from which to make the
http://dx.doi.org/10.1016/B978-0-12-394437-5.00004-8
 THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products 23

Figure 1 The grain-supply chain extends from the breeding of a variety to the product that reaches the consumer.

THE GRAIN CIRCLE countries, corresponding with breeders overseas and

The Consumer
Retail Breed
Process Sow
Store and
Grow
Transport
Harvest and
Segregate
Figure 2 The grain-supply chain can be visualized as a circle of
interactions and communications between the many functions.
selections. Over a century ago, entrepreneurial farmers selected
good-looking plants to keep as seed in hopes of plant
improvement.
The late nineteenth century brought the innovation of
crossbreeding and selection from the resulting progeny, with
the aim of combining the better attributes of each parent into
one variety. One of these early breeders was William Farrer,
represented in the drawings of Figure 3. Farrers vocation as a
surveyor in southeastern Australia led him to see that the wheat
varieties brought from England were unsuited to the new
country, so he set about importing seed from more suitable
crossbreeding.
Today, conventional crossbreeding involves the same basic
method as a century ago, namely, to take pollen from the
anthers of the male parent during anthesis and to dust
the pollen onto the stigma of the proposed female parent.
The anthers of the female have previously been removed to
prevent self-pollination, which is the usual form of reproduc-
tion for wheat and its close relatives. By contrast, the cross-
breeding of maize involves interfering with its normal
reproduction by which pollen, produced from the tassels on
top of the stalk, falls onto the silks protruding from the end of
the immature cob.
The tedious part of breeding is the subsequent task of
growing on the progeny and selecting desirable individuals
resulting from the cross. When elite lines have been selected,
they are grown on in field trials at various sites to see how they
perform in a range of conditions (Figure 4). After selecting the
most promising lines, they must be registered for plant
breeders rights, depending on the system operating in the
specific country, so that royalty payments may be returned to
the breeding organization.
The breeders involvement is primarily limited to introduc-
ing desirable genetic traits, but these benefits may subsequently
be modified by environmental factors during growth, such as
climate and farm management practices. Nevertheless, it has
proved possible for breeders to build in genetic tolerance to
some environmental risk factors, such as disease attack and
24 THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products

Figure 3 Depiction of a breeder at work in the 1890s when crossbreeding was in its infancy. (a) Cross-pollinating. (b) Examining progeny from a cross.
Note the great height of mature wheat plants of that time.

Figure 5 Equipment for sowing seed, together with fertilizer. In this

Figure 4 Breeders advanced lines, sown in relatively small plots for
further evaluation.
tolerance to premature sprouting in the case of a wet harvest.
Recent decades have seen the introduction of novel methods of
genetic manipulation, permitting breeders to employ a wider
range of options for genotype improvement.
Sow: Seed Must Be Clean and Genetically Pure
A critical decision for the grain grower is the choice of variety
for sowing, plus when to sow and in what situations on the
farm (Figure 5). The variety must suit the expected growth
case, it is designed for sowing narrow plots for breeder use.
conditions and the risks of disease and of environmental fluc-
tuations. Ideally, the grower attempts to target desired grain
qualities that would achieve premium prices upon delivery to
the mill or grain elevator. In addition, the grower must ensure
that the seed for sowing is free of weed seeds that would stifle
the desired crop, as well as causing the harvested grain to be
downgraded upon delivery due to the presence of contaminat-
ing seeds.
The source of seed may be saved-seed from a previous
harvest, or it may be obtained from a neighbor. However,
growers would ideally obtain pure seed (annually or every
few years) from a licensed pure-seed grower. Seed growers
 THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products 25

interact with breeders to obtain authentic seed of specific vari- When grain is delivered to the mill, storage silo, or grain
eties for growing under controlled conditions to ensure that elevator, it must be assessed for its quality and variety. This
the resulting grain conforms to the requirements for founda- assessment determines both the price to be paid to the grower
tion or registered seed. and the manner of segregation of the grain delivered. At any
delivery site, there may be several different grades or types of
Grow: Farm Management Must Optimize Grain Yield grain. It is obviously critical that a new delivery of grain should
and Quality be combined with grain of similar quality type in a distinct
storage cell.
In addition to decisions relating to seed for sowing, the grain
grower has decisions about soil preparation, fertilizer use at
sowing and during growth, sowing rate, and the possible use of
irrigation and of herbicides and pesticides. The task of maxi-
mizing financial returns involves on-farm management of
grain quality as well as grain yield, paying attention to the
combination of grain volume harvested, and market value
upon delivery. Even before harvest, the grower may have nego-
tiated marketing of the crop, based on expectations of yield
and quality attributes.

Harvest, Deliver, and Segregate: Grain Production Goes

Beyond the Farm Gate
Harvest is a busy time, with a sense of urgency to get the grain
into storage or delivered before rain comes. A wet harvest
brings the risk of the grain starting to sprout in the head, thus
being downgraded in price. Furthermore, most regions have an
upper limit on grain moisture, so the grain grower must ensure
that the grain is fully dried before harvesting the crop. Many Figure 7 A truck full of grain is being sampled from the elevated
large farms have storage facilities (Figure 6) so that the har- platform of the testing room, where the sample is evaluated to determine
vested grain can be stored immediately and the grower can its quality and thus where the load will be tipped.
then take some time about negotiating a good price for the
grain and appropriate conditions of transport.
Alternatively, as soon as the grain comes out of the header,
the harvested grain is trucked to a storage facility (elevator or
silo) or directly to a flour mill or feed mill (Figures 7 and 8).
Whereas bulk transport and storage are the general modes
today in most grain-growing areas, many years ago, grain was
packed in bags (Figure 9), which were man-handled and
stored in stacks. Regional centers for grain delivery and storage
are traditionally sited on rail lines because of the efficiency of
rail to transport large amounts of bulk grain (Figure 10(a)
and 10(b)).

Figure 6 A healthy field of awned wheat. After harvest, much of the

grain will be stored on farm in these bins, providing the grower with Figure 8 Grain being tipped from the farmers truck for delivery at the
flexibility to seek the best prices. country storage facility.
26 THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products
Storage and Transport: Identity Preservation Is Needed
for Grain Type and Quality
Further grain transport may involve delivery, probably by rail,
to the export terminal for loading into the hold of a ship
(Figure 11). In any case, some grain is likely to be stored on-
farm for seed use in the next season and for animal feed.
In situations of subsistence farming, the family will be directly
dependent on the stored grain for food throughout the
coming year.
Beyond the initial stage of storage, the value of the segre-
gated grain must be preserved throughout transport and on
through further storage so that it still conforms to the original
quality type when the grain is delivered for ultimate
processing. Throughout these stages, the grain is at risk of
deterioration especially due to insect attack. Also, throughout
these stages, commencing even before the farm gate, there is
the marketing of the grain at the level of the grain grower, by
grain merchants, and internationally often at inter-
governmental levels.
Figure 9 Bags of grain, as used for storage and transport prior to bulk
storage.
Figure 10 Grain storage facilities are generally situated beside rail connections to facilitate transport to the mill or export terminal.
Process: Production of the Many Grain-Based Foods
and Nonfood Items
It is at the stage of processing that the effectiveness of quality-
based segregation is evident and its cost-effectiveness can be
evaluated; for any situation, the advantages of better processing
outcomes must justify the significant costs involved in the
activities of testing and identity preservation. Due to these
commitments, grain-quality assessment is again a priority
when the grain arrives at the flour mill, malt house, feed mill,
or other forms of processing factory. This type of assessment
determines how a specific grain shipment will be stored,
utilized, and managed in the sequence of events leading to
the product that is offered for final sale to the consumer.
This sequence may be short: for example, the milling and
boiling of rice grown by a subsistence farmer, with the family
commenting on the eating quality. Even in a commercial situa-
tion, processing may be minimal involving whole-grain foods
such as rice, couscous, sweet corn, and beans (Figure 12). Min-
imal processing may involve sprouting to provide salad foods
(Figure 13). It may involve the stages of flour milling, leading to
bread and cake baking for sale in a hot-bread shop (Figure 14).
Further processes include the manufacture of breakfast cereals,
the use of cooked rice for sushi (Figure 15), the recovery of oils
Figure 11 Loading grain into the hold of a ship for international trade.
 THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products
Figure 12 Beans of various types, ready for sale without further
processing.
Figure 13 Sprouted beans. One more step of processing than in
Figure 12.
and their further processing, the separation of gluten from
wheat starch for paper manufacture, and the production of
sweeteners. Alternatively, there may be the much longer route
of animal feeding to produce meat, milk, cheese, or eggs via
grain-based pellets for animals or poultry (Figure 16).
Retail: The Stage When We Are All Involved in Expressing
Our Preferences
When the great diversity of grain-based products reaches the
retail stage, the ultimate in value addition is reached, with only
27
Figure 14 Bread rolls require the further stages of flour milling and
baking.
Figure 15 Sushi represents further value addition, compared to boiled
rice.
the extension of restaurant service taking value addition any
further. At this stage, the consumer (all of us) casts the votes
that determine the future success of the many variations of
products votes that may ultimately determine the future of
one baker against another or of one brand of breakfast cereal
versus others.
The Consumer
The consumer is thus the ultimate arbiter of quality for grain-
based products by voting with buying power. The identity of
the consumer ranges from the family members of the subsis-
tence farmer to the household expressing preferences for foods
from either the local baker or the supermarket. Thence, prefer-
ences by the respective bakers for flour quality pass up the
feedback loop back to the miller and so on to have repercus-
sions further back up the grain-supply chain (Figures 1 and 2).
28 THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products
Figure 16 Feed pellets, made from partly ground mixed grains, are
destined for animal feed a further stage of food production beyond the
stages in Figure 1.
This feedback loop from the consumer to others up the
chain may be short (a subsistence farming community) or
long (involving great distances and long time periods). The
latter case relates to the extent of worldwide grain trade as
the source of the grain involved. The ultimate feedback is to
the grain breeder, for whom a change in breeding direction
involves the many years from an initial cross to the selection of
an elite line ready for registration and sale to growers. Coming
back around the grain circle (clockwise in Figure 2), there is
the further lag factor of the many months that grain may be in
storage and transport before it is processed. In addition, feed-
back may involve long distances, such as a flour miller in the
Middle East or a legume trader in India reporting grain defects
to grain traders or breeders in grain-producing countries such
as the Americas, Australia, or Europe.
Consumer preferences also relate to their perception of
nutritional quality. Government agencies publicize nutritional
guidelines that generally recommend the consumption of
grain-based foods as a major part of food intake. This emphasis
on grains in the diet provides a major boost for the grain-
growing and grain-processing industries of the world. More
specific recommendations, such as for greater levels of soluble
fiber, may well target the breeder as the ultimate source of
significant changes in grain nutrition. Consumer feedback
may also target the processor with demands relating to the
incorporation of specific additives in foods as well as targeting
government agencies about legislation and food labeling.
The Biochemical Grain Chain
In contrast to the supply-chain concepts of Figures 1 and 2,
there is the biochemical concept of Figure 17 that sets out the
events in the context of genome, proteome, and grain compo-
sition, for the events in the developing grain, and the many
changes that occur during processing prior to the product
GENOME
Genes (DNA)
TRANSCRIPTOME PROTEOME
mRNA of expressed genes Polypeptides
ENVIRONMENT FUNCTIONAL PROTEINS
Climate, nutrients
CO2 GRAIN COMPOSITION
PROCESSING
PRODUCTS
Consumer acceptance
Figure 17 The sequence of biochemical events that occur in the grain,
leading to the grain-based products that reach the consumer.
Human
Arabidopsis
Rice
Wheat
Cotton
Barley
Figure 18 Comparative sizes of the genomes of several plant species.
Illustration kindly supplied by Dr. Colin Cavanagh, CSIRO, Canberra,
Australia.
reaching its ultimate destination the consumer. The general
sequence in Figure 17 is true for all grains with respect to their
production and end-use quality. Central to the sequence is the
genome the full DNA complement of genes. The relative sizes
of several genomes are depicted in Figure 18. Outstandingly
large is the genome of wheat, partly because common (bread)
wheat is a hexaploid, having three subgenomes, giving it a
complex genotype (the genetic makeup of the plant).
The first opportunity to examine the effects of growth con-
ditions, according to the scheme in Figure 17, comes by the
analysis of the transcriptome (mRNA) the transcripts from
those genes that are active under the environmental conditions
at the specific stage of grain development. This is the first
indication of what genes are being expressed in the endosperm
and also of how much of each gene product will be produced.
The next biological step is the resulting synthesis of the
many polypeptides corresponding to the mRNA transcripts
the proteome. The growth, yield, and suitability of grain for
processing of a grain are determined by the interaction of its
genotype (variety) with the environment (G  E). Genes being
expressed in the transcriptome (mRNA) are the focus of
research attention as candidates for use as markers for specific
attributes, thus being candidates for marker-assisted selection
in breeding. The corresponding polypeptides in the proteome
may also serve this purpose if the proteome is used as the target
 THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products
for marker selection. In either of these cases, the breeder has
the satisfaction of knowing that expressed genes are being
targeted, rather than genes in the genome that may not even
be expressed.
The transcriptome and the proteome also offer the advan-
tage of reflecting the effects of the growth environment. In
recent decades, we have learned much more about genotype
relationships than about the effects of environmental factors,
the latter being more difficult to study. With respect to wheat-
dough quality, research has concentrated on the genes coding
for the polypeptides that cross-link by disulfide bonds to form
the polymers of glutenin protein, the critical part of dough.
These polypeptides (subunits) are classed as the high- and
low-molecular-weight subunits, HMW-GS and LMW-GS,
whose many alleles are coded at the Glu-1 and Glu-3 loci,
respectively, on the Group-1 chromosomes of wheat. Informa-
tion about these gluten genes is available for many wheat
varieties from around the world. This allele information can
be used to predict dough properties as dough strength (Rmax)
and extensibility (Ext) (both parameters from the Brabender
Extensigraph), based on the alleles for both HMW-GS and
LMW-GS, assuming normal grain-filling conditions and aver-
age protein content.
Although we realize that wheat quality is also governed by
some other protein factors, this information is valuable to
breeders and further down the grain chain for quality-based
segregation at harvest. However, there are still serious gaps in
our knowledge surrounding the effects of fluctuations in
growth conditions.
More important, but more poorly understood, is the man-
ner by which these glutenin polypeptides are disulfide cross-
linked into functional glutenin (Figure 17). The glutenin poly-
mers are so large that conventional methods of protein analysis
are inadequate. Field-flow fractionation (FFF) has been used to
examine the upper size ranges, which are critical to dough
strength. FFF has shown, for example, that the dough-
weakening effects of high temperatures during grain filling
are accompanied by a reduction in the proportion of very
large glutenin polymers.
However, we are yet to form a valid understanding of the
factors that determine the formation of disulfide bonds
between glutenin subunits and the extent of the ongoing poly-
merizations. Models have been suggested for the interactions
between the HMW-GS and LMW-GS and for polymeric glute-
nin with gliadin proteins, but we do not understand the G  E
aspects of this very significant stage of grain-quality
determination.
We are aware that the formation of disulfide bonds within
the gluten complex can continue during storage, especially at
temperatures over about 30  C. In addition, disulfide bonds
are a target for manipulation during processing (P) the third
factor that determines end-product quality (G  E  P).
Past the farm gate, there is a marketing opportunity that we
cannot presently grasp fully the possibility of segregating
grain deliveries based on their suitability for end-product man-
ufacture, considering the effects of both G and E. At this stage,
knowledge of variety is of less importance than of quality type,
which might be better specified by quantitative analysis of
glutenin alleles. Some promise is offered for this purpose by
on-the-spot equipment for the analysis of glutenin-subunit
29
composition to permit segregation prior to dumping
(comingling), after which distinct identity is lost. More specific
knowledge of the grain quality of specific storage cells offers
valuable market intelligence for use in maximizing return on
grain and in providing the processor (next in the grain chain)
with grain of the required quality specifications.
Future Prospects
The general sequence of grain production and processing in
Figures 1 and 2 must continue into the future, but there will be
changes within each of the stages. The breeding of new and
improved genotypes (varieties) is ongoing, so too are develop-
ments in agronomy and farm management. However, these
strategies will need to change and develop in view of the
ongoing climate challenges.
The rising levels of carbon dioxide in the atmosphere con-
stitute an effective carbon fertilizer. Field experiments with
ambient and elevated carbon dioxide have clearly shown that
the carbon fertilizer effect produces greater yields of biomass
and of grain. However, the protein content of the grain is
lower, thus affecting grain quality and utilization.
Grain production will also be affected by predictions of
rising temperatures, shifting rainfall patterns, and more fre-
quent stress situations such as heat shock, storms, and violent
winds. Furthermore, rising sea levels present the risk that some
agricultural regions will become unsuitable for cropping.
The grain-production industry worldwide needs to able to
adapt to a changing climate while also ensuring that the nec-
essary changes are consistent with sustainable production. In
line with these production challenges, markets will require
greater uniformity and certainty of grain-quality specifications,
although these discerning markets must be prepared to pay
premiums in contrast to commodity-level grain.
Processors and retailers will meet more customer-based
pressure to supply specific nutritional requirements, while
large supermarket companies will exert downward pressure
on suppliers.
Exercises for Revision
What features distinguish the added-value route of one
grain species from another? Compare the stages for two or
three contrasting grain species.
Describe some grain-based products that have shorter and
longer chains.
What are some critical feedback messages that are indicated
by the grain chains in Figures 1 and 2?
The shorthand expression G  E  P is used in relation to
Figure 17. To what extent does it explain the grain chain?
Exercises for Readers to Explore the Topic Further
Select some grain-based products and outline the grain
chain that has led to the production of each of these.
Some examples are
high-fructose syrups used as sweetening agents,
30 THE BASICS | The Grain Chain: The Route from Genes to Grain-Based Products
margarine from canola or any other oilseed,
cornflakes or another breakfast cereal,
ethanol-based biofuels,
oilseed-based diesel fuel.
In your region, identify the main grains that are grown and/
or processed. What forms of quality-based segregation,
storage, and transport are used?
Investigate the biochemical sequence in Figure 17 for some
grain species other than wheat.
Protein isolates are produced from several grains. Investi-
gate the production and uses of some, such as wheat gluten,
soy-protein isolate, and corn protein (sometimes called
corn gluten).
Examine some current consumer issues related to grain-
based foods. In which of the upstream parts of the grain
chain is there the need for more diligent attention to
quality?
See also: Barley, Rice and Maize Processing: Rice Processing:
Beyond the Farm Gate; Breeding of Grains: Maize: Breeding; Wheat
Breeding: Exploiting and Fixing Genetic Variation by Selection and
Evaluation; Genetics of Grains: Genomics of Food Grains; Grain
Harvest, Storage and Transport: Wheat: Harvesting, Transport,
and Storage; Grain Marketing and Grading: Barley: Grading and
Marketing; Wheat: Grading and Segregation; Non-wheat Foods:
Maize: Foods from Maize; Rice: Chinese Food Uses; Processing of
Grains: Grain Quality Attributes for Cereals Other than Wheat;
Evaluation of Wheat-Grain Quality Attributes; The Cereal Grains: An
Overview of the Family of Cereal Grains Prominent in World
Agriculture; Wheat: An Overview of the Grain That Provides Our Daily
Bread; The Legumes and Pseudocereals: Grain Legumes and
Their Dietary Impact: Overview; The Oilseeds: Oilseeds: Overview;
Wheat-Based Foods: Cereals: Overview of Uses: Accent on Wheat
Grain.
Further Reading
Abdel-Aal E and Peter Wood P (eds.) (2005) Specialty Grains for Food and Feed.
St Paul, MN: AACC International Press.
Arendt EK and Zannini E (eds.) (2013) Cereal Grains for the Food and Beverage
Industries. Oxford: Woodhead Publishing Ltd.
Faridi H and Faubion J (1995) Wheat End Uses Around the World. St. Paul, MN:
American Association of Cereal Chemists.
Henry RJ and Kettlewell PS (eds.) (1996) Cereal Grain Quality. London: Chapman and
Hall.
Kent NL and Evers AD (eds.) (1994) Technology of Cereals. An Introduction for
Students of Food Science and Agriculture, 4th edn. Oxford: Pergamon Press.
Wrigley CW and Batey IL (eds.) (2010) Cereal Grains: Assessing and Managing Quality.
Oxford: Woodhead Publishing Ltd.
 Taxonomic Classification of Grain Species
LA Morrison, Oregon State University, Corvallis, OR, USA
CW Wrigley, The University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Modern taxonomy is a uniform system for assigning scien-
tific names to plant species, thereby aiding communication
among scientists.
Classification is structured as a hierarchical system that
follows a concise set of rules for both wild and domesti-
cated plant species.
Species names often convey information about a grains
function or traits or the activities associated with its use.
Grains can be divided into two large groups, each with
different evolutionary histories: cereals (grass species) and
pseudocereals and pulses (nongrass species categories).
Taxonomic groupings reflect evolutionary relationships
with interpretations often varying among traditional versus
phylogenetic classifications.
For domesticated plants, the terms cultivar and landrace
are not interchangeable.
Taxonomic classification enables informed decisions for
accurately identifying and selecting research material of
grain species.
Evolutionary relationships are useful for identifying poten-
tial allergic cross-reactions to grain species.
Using classification to display phylogenetic relationships is
at the center of current controversies over the future direc-
tion of Linnaean taxonomy.
Learning Objective
To achieve understanding of the role of taxonomy as a scien-
tific system for both classifying domesticated grain species
and facilitating communication about them among scientists
Introduction
The common names of grain species often derive from local-
ized, figurative terms. They can lead to considerable confusion
due to variations in region and language. In English, the prob-
lem is exacerbated by the adoption of common names from
several languages, either as Anglicized words or direct trans-
lations. The grain legume Cicer arietinum offers a very good
example of multiple common names. In languages of Europe
and Asia, it is known as cece (Italian), channa dhal (Hindi),
garbanzo (Spanish), grao-de-bico (Portuguese: beaked
grain), hummus (Arabic), Kichererbse (German: giggle
peas), and pois chiche (French: chickpea). In English, it is
variously known as chickpea, gram, Bengal gram, chana, Egyp-
tian pea, and garbanzo bean. Another problem is the changed
usage of a common name from one region to another. In
Europe, corn refers to cereal grains generally, whereas in the
Americas, it specifically refers to maize (Zea mays).
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00002-4
Taxonomy: A System of Names to Avoid Confusion
Common names can often have multiple or misleading asso-
ciations. If not qualified, pigweed can collectively refer to
members of the genus Amaranthus without distinguishing
between the weedy and grain amaranth forms. The wheat in
buckwheat (Fagopyrum esculentum; Figure 1) and Inca wheat
(Amaranthus caudatus; Figure 2) creates the false impression of
a relationship with wheat (Triticum species; Figure 3), when in
fact wheat and these two pseudocereal species are far apart
from each other in evolutionary relationships, in biology,
and in grain attributes. A more reliable means of communicat-
ing is provided by scientific names, which are assigned by a
system known as taxonomy.
A scientific or taxonomic name consists of a genus and a
species name that, when combined as a binomial, provide a
unique identifier for the grain in question. This name is uni-
versally recognized in all scientific circles and removes the
confusion created by common names, as the following exam-
ple illustrates. Amaranthus caudatus, A. creuntas, and A. hypo-
chondriacus form the group known as grain amaranths, major
food staples in the pre-Columbian Americas, which have now
regained importance and promise as pseudocereal grains in
international agriculture. Their taxonomic names distinguish
these grain amaranths from other weedy Amaranthus species
variously known as pigweed, amaranth, or water hemp.
Although some common names have been accepted into
international usage, they can still create problems within a
scientific context. For example, in large species groups such as
the wheats, scientific names make distinctions where common
names cannot. The term wheat does not differentiate between
the wild and domesticated species of Triticum nor between the
species of Triticum and its sister genus Aegilops, whose members
are all wild species. The common name farro can collectively
refer to any one of three domesticated wheat species, all hulled
(non-free-threshing) T. monococcum, a diploid wheat that is a
minor crop species; T. dicoccum, a primitive tetraploid wheat
also known as emmer; and T. spelta (T. aestivum ssp. spelta), a
hexaploid wheat known as spelt that is still grown in Europe and
Asia and now a specialty wheat in the United States and Canada.
Common wheat and bread wheat are equivalent names that can
encompass a complex of domesticated hexaploid wheats when
they are classified under one species name, Triticum aestivum,
thereby including forms that are neither common nor necessar-
ily amenable to bread-making (Figure 3).
Taxonomy: A System of Relationships
Inherent in taxonomic groupings is an indication of evolution-
ary relationships. Thus, the taxonomy of grain species has
some bearing on shared traits of plant morphology, grain
31
32 THE BASICS | Taxonomic Classification of Grain Species
Figure 1 Buckwheat (Fagopyrum esculentum). Its dried, triangular-
shaped fruit, a nut, resembles the beechnut produced by beech trees
(Fagus spp.). This similarity is the source for the genus name
Fagopyrum, which translates to beechwheat. It also explains the
derivation of the English common name buckwheat. Buck is the
Anglicized version of the German word for beech, Buche. Adapted from
Williams JT (ed.) (1995) Cereals and Pseudocereals. London: Chapman
and Hall.
composition and function, and many other attributes impor-
tant to grain science. As a tool for categorizing biological
organisms, taxonomy is not an exact science and thus is subject
to conceptual disagreements and opposing interpretations.
Current debates primarily center on its ability to accurately
represent phylogeny the branching ancestor/descendant rela-
tionships that connect all plant species. On one side are molec-
ular systematists, who argue that classifications must reflect
evolutionary relationships. On the other side are traditional-
ists, who counter that the reliable order and structure of tax-
onomy is more important than an exact representation of
modern phylogenetic concepts that are often changing in
response to new research evidence. Problematic for phyloge-
netic classifications are (1) their continuous state of flux based
Figure 2 A mature specimen of Amaranthus caudatus, the most
important of the Andean grain amaranths. It is still known by the ancient
Incan name kiwicha. Seeds (left) are about 1 mm in diameter and have
a curved embryo. Reproduced from Vietmeyer ND (ed.) (1989) Lost Crops
of the Incas. Washington, DC: National Academy Press, with permission.
on the availability of new genetic evidence supporting revi-
sions of accepted and unresolved groups and (2) their inability
to accommodate interspecific hybridization and introgression,
two plant speciation mechanisms that violate the monophy-
letic rule that all descendants in a phylogenetic tree derive from
a single common ancestor species. Another key element in
taxonomic debates is how to construct uniformly accepted
treatments, particularly for domesticated species. Grain
researchers often encounter conflicting views in differing
names and classification schemes for the same grain species
or species groups. In the midst of such debates, the taxonomic
system developed in the eighteenth century by the Swedish
scientist Carl Linnaeus is still intact. While providing a rela-
tively flexible structure for building modern classifications, the
Linnaean traditional approach faces challenges when robust
genetic evidence reveals classification inconsistencies with a
groups actual evolutionary relationships. For additional detail
on the practice of taxonomy and classification issues related to
evolutionary relationships.
 THE BASICS | Taxonomic Classification of Grain Species 33

Figure 3 Variations in the appearance of heads of wheat species, one of many morphological characteristics used for their taxonomic classification.
From left to right, the wheat species are (including their genome assignments and common names): T. boeoticum (2x: wild einkorn), T. monococcum
(2x: einkorn), T. dicoccoides (4x: wild emmer), T. dicoccum (4x: emmer), T. durum (4x: macaroni wheat), T. carthlicum (4x: Persian wheat),
T. turgidum (4x: rivet wheat, cone wheat), T. polonicum (4x: Polish wheat), T. timopheevii (4x: Timopheevs wheat), T. aestivum (6x: bread wheat),
T. sphaerococcum (6x: shot wheat, Indian dwarf wheat), T. compactum (6x: club wheat), T. spelta (6x: spelt), and T. macha (6x: macha wheat). The
diploid A-genome species, T. urartu, is not shown here. 2x diploid; 4x tetraploid; 6x hexaploid. Adapted from Mangelsdorf PC (1953) Wheat.
Scientific American 189: 5059, 1953 by Scientific American Inc.

Taxonomy: What It Is and How It Started
Taxonomy provides a system by which plants are classified,
named, and identified. Classification consists of the circum-
scription, grouping, and hierarchical ranking of a plant entity
(taxon) or entities (taxa). Naming, or nomenclature, is gov-
erned by a set of rules laid out in the International Code of
Botanical Nomenclature (ICBN) and by the specialized rules
for domesticated plants found in the International Code of
Nomenclature for Cultivated Plants (ICNCP). Identification,
34 THE BASICS | Taxonomic Classification of Grain Species
which usually goes hand in hand with nomenclature, is a
determination based on similarity of a taxons membership
in an existing group or a new group.
Linnaeus initiated modern taxonomy with the publication
of Species Plantarum in 1753, which classified every plant then
known to him. The Linnaean system was not an isolated
endeavor. It was built on a long history of scientific attempts,
beginning with the Greek Theophrastus (third century BC) and
the Roman Pliny the Elder (first century AD), to construct
universal naming and classification systems. The Linnaean
binomial system, consisting of genus and species names,
resolved the problem of concisely and uniquely naming a
plant. Continuing with the scientific heritage that preceded
him, Linnaeus used Latin as the language of taxonomic
names and descriptions. In his time, Latin was the universal
language by which European scholars could communicate.
Linnaeus took classical Latin and created a new botanical
form with its own terminology for plant structures and the
flexibility for adopting Latinized words taken from other lan-
guages. Modern attempts to replace botanical Latin with
English for taxonomic descriptions have so far been unsuccess-
ful, leaving this particular Linnaean heritage intact.
Origins of Taxonomic Names
Many Linnaean scientific names for agricultural plants have
their roots in the vernacular of the ancient Greeks and Romans.
For example, the genus names for grains were derived from
terms that often related to a particular function, activity, or trait
or to an association with a person. Thus, Triticum comes from
the Latin tero meaning I thresh; Cicer, which translates to
chickpea, likely has its origin in the Greek kikus meaning
strength and power, a feature of chickpeas embodied in their
high nutritional value and a reputation as an aphrodisiac. For
Cicer, the story is also tied up in family names. In time, the
early Greek name for chickpea evolved to kiros, which became
cicer in Latin. Early Roman families would take surnames that
related to their prosperity. Given the importance of legumes, it
is not surprising that Cicero was the family name taken from
cicer; likewise three other famous Roman families carried
legume names: Fabius (faba: fava bean), Lentulus (lens: len-
til), and Piso (pisum: pea).
In the original Linnaean system, species names usually
conveyed information about a trait. For example, cereale traces
to Ceres the Roman goddess of agriculture and more specifi-
cally to Cerealis (of or relating to Ceres) from which the term
cereals was derived for collectively naming the edible grains of
grass species. The genus and species names together produce an
identifying designation for taxa as these examples illustrate:
Amaranthus caudatus translates as the amaranth ending with a
taillike appendage, a reference to this species long, taillike
inflorescence (Figure 2); Cicer arietinum as chickpea shaped
like a rams head, a reference to its seed shape; Fagopyrum
esculentum as edible buckwheat (Figure 1); Triticum monococ-
cum as one-grained wheat (Figure 3); and Secale cereale as
cereal rye.
For species named by Linnaeus, the intertwining of scien-
tific names with the original vernacular helps to explain their
respective derivations. The origin of the term common wheat
hails back to an older, no longer accepted, species name
T. vulgare, which translates as common wheat. The use and
derivation of corn and the genus name, Zea, provide another
example. The English word corn has its roots in the German
word for grain, korn. The Greek word zea, like corn, referred
to grain in a general sense. Linnaeus selected Zea as a genus
name for the new cereal grain species that had arrived in
Europe following the discovery of America and for which
there was no European vernacular name upon which to draw.
As a consequence, the English common name corn was
adopted into the American English vernacular to refer to the
only major New World cereal grain known in the eighteenth
century.
The Use of Genus and Species Names
Tables 1 and 2 list common and scientific names for grain
species. The convention is to write both genus and species
names in italics, with the genus name having a capital as the
first letter, followed by the species name (known as the specific
epithet) all in lower case. When a grain consists of a group of
species, the plural abbreviation spp. is used. In text, the genus
name component of the binomial is spelled out the first time
and abbreviated to its first letter, except when beginning a
sentence, for each following use as was done earlier in the
text with the species examples of Amaranthus and Triticum. In
cases where two genera with the same first letter are discussed
together, then the second letter of the species name is included
in the abbreviation, for example, in discussions of the millets,
Pa. for Panicum and Pe. for Pennisetum.
Following the species and subspecies epithets is the name
(usually abbreviated and not in italics) of the botanist or
researcher who named the species, for example, L. for Lin-
naeus in T. monococcum L. and Desf. for R. L. Desfontaines in
T. durum Desf. Known as the authority, this part of the scien-
tific name is important for synonymy, the component of tax-
onomy that deals with the historical naming of a species. When
the name and classification of a grain species are in question
due to competing treatments, synonymy provides a way to
trace its taxonomic handling and to determine which is the
correct and legitimate name according to the ICBN. Authority
names are not included in Tables 1 and 2 as they are not
essential for the completeness of the binomial in text discus-
sions such as this. However, it is useful in research articles and
literature reports to include the authority citation at the first
mention of the species to clarify which taxonomic concept is
being followed.
Taxonomy: Rank and Hierarchy
In classification, taxonomy provides a system for grouping
species into ranked hierarchies. All grain species are flowering
plants and thus are classified together as angiosperms. Mem-
bers of this group reproduce sexually with the male and female
organs located in the flowers, which can be showy (pseudocer-
eals, pulses, and oilseeds) or inconspicuous (cereals). Their
seeds are protected by a fruit, which can be fleshy or dried. In
the case of cereals, the fruit is reduced to a thin outer tissue that
 THE BASICS | Taxonomic Classification of Grain Species 35

Table 1 Classification of the grass family, Poaceae (Gramineae), all monocotyledonous plants: cereal grain species and their common namesa

Subfamily Tribe Genus and species Common name

b
Ehrhartoideae Oryzeae Oryza glaberrima African rice
Oryza sativa Rice
Zizania aquatica Wild rice
Zizania palustris Northern wild rice
Chloridoideae Eragrostideae Eleusine coracana Finger millet, African millet, Indian millet
Eragrostis tef Tef, teff, teff grass
Panicoideae Andropogoneae Sorghum bicolor Sorghum, grain sorghum, great millet, Kaffir corn, durra, milo
Zea mays Corn, maize
Coix lacryma-jobi Coix, Jobs tears, Adlay millet
Paniceae Digitaria exilis Fonio, fonio millet, hungry rice
Echinochloa esculenta Japanese barnyard millet, Japanese millet, Siberian, or marsh millet
Panicum miliaceum Common millet, broom millet, broomcorn millet, proso millet
Panicum sumatrense Blue panic, little millet, sama
Paspalum scrobiculatum Kodo millet
Pennisetum glaucum Pearl millet, cattail millet
Setaria italica Foxtail millet, Italian millet
Pooideae Poeaeb Avena sativa Oat, side oat
Avena abyssinica Abyssinian oat
Phalaris aquatica Bulbous canary grass, Harding grass, Toowoomba canary grass
Triticeae Hordeum vulgare Barley
Secale cereale Rye, cereal rye
Triticosecalec Triticale
Triticum aestivum Wheat, common wheat, bread wheat
Triticum dicoccum Emmer, farro
Triticum durum Durum wheat, hard wheat, macaroni wheat
Triticum monococcum Einkorn, small spelt, farro
Triticum spelta Spelt, spelt wheat, hulled wheat, dinkel wheat, farro
Triticum turgidum Rivet wheat, cone wheat
a
Following the classification system outlined in Heywood VH, Brummitt RK, Culham A, and Seberg O (2007) Flowering Plants of the World. Ontario, Canada: Firefly Books, and
updated by the Angiosperm Phylogeny Group.
b
Subfamily and tribal classification assignment according to the Grass Phylogeny Working Group (GPWG).
c
Triticale (Triticosecale) is an intergeneric hybrid that was created by crosses between wheat and cereal rye. The  preceding Triticosecale, which combines Triticum and Secale,
indicates its hybrid origin. Given the multiple hybrid combinations by which Triticale forms have been derived, only cultivar names are assigned.

is adherent to the seed. In botanical terminology, the cereal
grain is called a caryopsis.
Traditional Linnaean classifications have followed the hier-
archical structure of KingdomClassOrderFamilyTribe
GenusSpecies with subdivisions in each of these ranks, for
example, subclass. Largely morphological in concept, these
classifications do not always track well with current evidence
on evolutionary relationships. On the other hand, phyloge-
netic classifications are structured around monophyletic
groups that each consist of all descendants of a common
ancestor, which share the groups identifying characteristics.
Given that ongoing molecular genetic research leads to chang-
ing evolutionary concepts, phylogenetic classifications are in
flux and have unresolved groupings. While some of the lower
Linnaean hierarchical rankings have been retained in these
classifications, groupings above Order, as well as unresolved
groups at lower ranks, are designated as clades. As a general
term denoting a group consisting of an ancestor and all its
descendants, clade is used in this latter context to indicate a
group whose phylogenetic status is still open to question.
Differences distinguishing traditional versus phylogenetic
classifications come into play for the monocot and dicot evolu-
tionary lines of grain species. In older classifications, flowering
plants belong to the Class Angiospermae, which is divided into
monocots (subclass Monocotyledoneae) and dicots (subclass
Dicotyledoneae). From a more current phylogenetic perspective,
angiosperms are classified as a clade with three subgroupings:
dicots, monocots, and eudicots. Monocots and eudicots (true
dicots) are each designated as a clade (Table 3). Dicots are not
treated as a clade because they are a nonmonophyletic group of
primitive angiosperms with various members ancestral to both
monocots and eudicots, respectively.
Grain species covered in this article fall within the monocot
and eudicot clades. Therefore, the biological features associated
with monocots and dicots are useful for distinguishing among
cereal (monocot) and noncereal (dicot) species. The terms
monocot and dicot refer to the presence of one or two embryonic
leaves (cotyledons) in the seed and young seedling. Other easily
identifiable morphological traits differentiate between them.
Monocots have flower parts in threes, parallel-veined leaves,
scattered vascular bundles, and fibrous adventitious root system.
Dicots have flower parts in fours or fives, net-veined leaves,
vascular bundles located in a ring, and a primary tap root.
Tables 13 present current taxonomic treatments for cereal
and noncereal grain and oilseed species. For additional detail
on traditional and phylogenetic classification approaches.
Family is the most useful of the higher taxonomic ranks.
Family names end in -aceae except for several older names
36 THE BASICS | Taxonomic Classification of Grain Species

Table 2 Dicotyledonous grain and oilseed species and their common namesa

Family Tribe Genus and species Common name

Amaranthaceae Amaranthoideae Amaranthus caudatus Amaranth, grain amaranth, Inca wheat, kiwicha
Amaranthus cruentas Purple amaranth, red amaranth
Amaranthus hypochondriacus Princes feather
Atripliceae Chenopodium quinoa Quinoa
Asteraceae (Compositae)b Cynareae Carthamus tinctorius Safflower
Heliantheae Guizotia abyssinica Niger, niger seed, ramtil
Helianthus annuus Sunflower
Brassicaceae (Cruciferae)b Brassiceae Brassica napus Canola, oilseed rape, rape
Brassica spp. Mustard
Crambe abyssinica Crambe
Fabaceae (Leguminosae)b Dalbergieae Arachis hypogaea Peanut, groundnut, goober
Cicereae Cicer arietinum Chickpea, garbanzo bean, gram
Genisteae Lupinus albus White lupin
Lupinus angustifolius Blue lupin
Phaseoleae Cajanus cajan Pigeon pea, Congo pea, red gram
Glycine max Soybean
Phaseolus lunatus Lima bean, butter bean
Phaseolus vulgaris Bean, kidney bean, pinto bean, navy bean, cannellini bean
Vigna angularis Adzuki bean, azuki bean, aduki bean
Vigna mungo Black gram, urad bean
Vigna radiata Mung bean, golden gram, green gram
Vigna unguiculata Black-eyed pea, cowpea
Fabeae Lathyrus sativus Chickling pea, chickling vetch
Lens esculentus Lentil
Pisum sativa Garden pea, field pea
Vicia faba Faba bean, broad bean
Lamiaceae (Labiatae)b Mentheae Salvia hispanica Chia, salba
Linaceae Linum usitatissimum Linseed, flax, Linola
Malvaceae Gossypieae Gossypium spp. Cottonseed
Pedaliaceae Sesamum indicum Sesame, sesame seed, benne, benne seed
Polygonaceae Fagopyrum esculentum Buckwheat, Japanese buckwheat, silverhull buckwheat
a
Following the classification system to family rank outlined in Heywood VH, Brummitt RK, Culham A, and Seberg O (2007) Flowering Plants of the World. Ontario, Canada: Firefly
Books and updated by the Angiosperm Phylogeny Group.
b
Original family name predating ICBN standardization.

that were in use before the ICBN standardized the suffix spell-
ing. Thus, in grain taxonomy, two family names are occasion-
ally encountered Gramineae (old) and Poaceae (new),
Leguminosae (old) and Fabaceae (new), Cruciferae (old) and
Brassicaceae (new), and Compositae (old) and Asteraceae
(new). The other useful higher taxonomic ranking is tribe, a
subdivision of family, that is identified by an -eae ending. All
the cereal grains (e.g., maize, rice, sorghum, and wheat) are
monocots and belong to the grass family Poaceae. Dicot grains
and oilseeds comprise a diverse mixture of families. The pulses
or grain legumes (e.g., beans, peas, and soybeans), which are
the largest group of dicot grains, are classified in Fabaceae.
Other families include one or just a few species, for example,
Amaranthaceae (grain amaranths and quinoa), Polygonaceae
(buckwheat), Asteraceae (niger, safflower, and sunflower),
Malvaceae (cottonseed), and Brassicaceae (canola, mustard,
and crambe). Subdivisions into subfamilies and/or tribes
reflect the size and diversity of the family.
Grain species can be subdivided into infraspecific categories
known as subspecies (ssp.), variety (var.), and forma (f.).
Because there is no clear agreement on use of subspecies versus
variety, the two are often used interchangeably, although it is
possible to treat variety as a subrank to subspecies. When the
subspecies (or variety) is the taxonomic type that defines a
particular species, then the ICBN requires that this subspecies
(or variety) repeats the species name for example, T. aestivum
ssp. aestivum. In general discussions, it is not necessary to
include the repetitious infraspecific form. Thus, in this exam-
ple, it is sufficient to cite only the species name T. aestivum
when referring to bread wheat. Forma is the smallest category
and is infrequently encountered.
The infraspecific ranks are intended to describe variants that
form distinct groups defined by genetics, morphology, ecology,
and distribution. How a particular taxonomist chooses to clas-
sify infraspecific taxa can appear arbitrary. For example, a current
taxonomic treatment of the wheats uses variety for the wild
wheat genus Aegilops (e.g., A. speltoides var. ligustica) but subspe-
cies for the domesticated genus Triticum (e.g., T. turgidum ssp.
durum). Here, the infraspecific ranks should be viewed as
approximately equivalent. This example also illustrates the dif-
ficulty in reflecting evolutionary relationships within the frame-
work of taxonomic classification. The infraspecific variation in
domesticated Triticum species is the product of human selection
pressure, which is arguably on a different evolutionary scale
than the variation in the wild species of Aegilops. Thus, in this
example, the taxonomists choice of variety for Aegilops and
subspecies for Triticum should not be construed as a definitive
reflection on evolutionary status.
 THE BASICS | Taxonomic Classification of Grain Species 37

Table 3 Hierarchical classificationa for grain and oilseed speciesb with their use-category assignments

Cladec Order Family Subfamily Tribe Genus Use category

Monocots
Commelinids Poales Poaceae Ehrhartoideaed Oryzeae Oryza Cereal
Zizania Cereal
Chloridoideae Eragrostideae Eleusine Cereal
Eragrostis Cereal
Panicoideae Andropogoneae Coix Cereal
Sorghum Cereal
Zea Cereal
Paniceae Digitaria Cereal
Echinochloa Cereal
Panicum Cereal
Paspalum Cereal
Pennisetum Cereal
Setaria Cereal
Pooideae Poeaed Avena Cereal
Phalaris Cereal
Triticeae Hordeum Cereal
Secale Cereal
Triticosecale Cereal
Triticum Cereal
Eudicots
Core eudicots Caryophyllales Amaranthaceae Amaranthoideae Amaranthus Pseudocereal
Chenopodioideae Atripliceae Chenopodium Pseudocereal
Polygonaceae Fagopyrum Pseudocereal
Asterids Asterales Asteraceae Lactucoideae Cardueae Carthamus Oilseed
Asteroideae Heliantheae Guizotia Oilseed
Helianthus Oilseed
Lamiales Lamiaceae Nepetoideae Mentheae Salvia Oilseed
Pedaliaceae Sesamum Oilseed
Rosids Fabales Fabaceae Papilionoideae Aeschynomeneae Arachis Pulse, oilseed
Cicereae Cicer Pulse
Genisteae Lupinus Pulse
Phaseoleae Cajanus Pulse
Glycine Pulse, oilseed
Phaseolus Pulse
Vigna Pulse
Vicieae Lathyrus Pulse
Lens Pulse
Pisum Pulse
Vicia Pulse
Geraniales Brassicaceae Brassiceae Brassica Oilseed
Crambe Oilseed
Malpighiales Linaceae Linoideae Linum Oilseed
Malvales Malvaceae Malvoideae Gossypieae Gossypium Oilseed
a
Following the classification system to family rank outlined in Heywood VH, Brummitt RK, Culham A, and Seberg O (2007) Flowering Plants of the World. Ontario, Canada: Firefly
Books as updated by the Angiosperm Phylogeny Group.
b
For additional information on individual taxa.
c
To view the associated phylogenetic trees and clade subdivisions, see http://www.mobot.org/MOBOT/research/APweb/welcome.html.
d
Alternative subfamily and tribal classification assignment following the Grass Phylogeny Working Group (GPWG) revision.

Taxonomic Disagreements
With taxonomic classification, there is not necessarily a right or
a wrong treatment. Taxonomists and the researchers working
with a particular group can often disagree with each other. As a
consequence, different names for the same species can be
found in use. For those who are not schooled in the ICBN or
unfamiliar with the underlying details of a taxonomic contro-
versy, conflicting classifications are confusing and can be
difficult to sort out. In the case of domesticated species, debates
usually deal with both the taxonomic status of taxa (usually
species versus an infraspecific rank) and the relative impor-
tance of genetic evidence in defining species concepts and
evolutionary relationships. There is no clear separation of
these two points, partly due to the historical tradition on
which taxonomy has been built and the inconsistencies in
how domesticated plants are viewed within an evolutionary
context.
38 THE BASICS | Taxonomic Classification of Grain Species
The opposing species concepts at play in wheat taxonomy
offer an example of the different directions that a taxonomic
debate can take. Morphological diversity in the domesticated
tetraploid and hexaploid wheats illustrates the potential for
variation that developed with humans acting as evolutionary
selection agents. This variation originally correlated with eco-
geographic distributions and, if allowed to continue in isolation
over geologic time, would have possibly led to new species.
Wheat geneticists stress the need to portray accurate genetic
relationships, circumscribing wheat species on the basis of
their close genetic similarities. Thus, in the AB- and ABD-
genome groups, geneticists accept only two species, each with
several infraspecific forms T. turgidum and T. aestivum. On the
other side of the fence are those who stress the importance of
recognizing the distinct variation among domesticated forms by
naming multiple tetraploid and hexaploid wheat species
(Figure 3 and Table 1). Provided that the rules of nomenclature
are followed, differing treatment concepts are acceptable. On
this basis, T. spelta and T. aestivum ssp. spelta are both correct as
are T. durum and T. turgidum ssp. durum, respectively, describing
the same wheats, albeit from different taxonomic perspectives.
Cultivated Plant Taxonomy
Grain and oilseed species are classified as botanical entities
under the ICBN. Their man-made forms derived through arti-
ficial selection, known by the general term cultigens, are clas-
sified under the ICNCP in two possible categories: cultivar
(deliberately selected plants, either domesticated or wild) and
group (an assemblage of cultivars). The term cultivar was
developed from cultivated variety by combining the first five
and three letters of these two terms, respectively. According to
the ICNCP, a cultivar is designated by one or more distinct,
uniform, and stable characters that persist through propaga-
tion. Although often used interchangeably, the terms cultivar
and variety are not equivalent, that is, variety is not a nomen-
clatural term under the ICNCP. However, in the context of
cultivar registration, the term variety is often used to refer to a
cultivar. Of note, variety is a legal term under the Interna-
tional Union for the Protection of New Varieties of Plants
convention, which oversees the recognition of plant breeders
rights for a particular domesticated cultivar. Regarding termi-
nology differences between the ICBN and the ICNCP, the
botanical term variety (also as varietas) applies only as a
name for one of the infraspecific ranks under the ICBN.
A domesticated cultivar has a non-Latin name (cultigen
epithet) that is not italicized and enclosed by single quotation
marks. Several examples should help to clarify this usage Zea
mays Bronze Beauty or maize cultivar Bronze Beauty, Lens
culinaris Laird, Avena sativa Wallaroo, Arachis hypogaea ssp.
fastigiata Georgia Red, Amaranthus cruentus K343, and Cicer
arietinum Kranthi. Following a cultivars first mention, the
single quotation marks can be dropped.
The term landrace is often encountered in discussions of
crop germplasm. It refers to an indigenous grain species within
a local farming system. Landraces predate modern agriculture
and often are ancient infraspecific forms that may have been in
cultivation for hundreds or thousands of years. Landraces are
usually composed of a mixed population whose members vary
by agronomic traits such as disease resistance, maturity, yield,
and cold tolerance. The mixture also may include different
species. For example, wheat landraces collected during the late
1940s in southeastern Turkey can be composed of ten or more
different forms of the two species T. aestivum (T. aestivum ssp.
aestivum) and T. durum (T. turgidum ssp. durum). The advantage
of landraces in subsistence farming systems is the guarantee of a
crop under all types of growing conditions. For example, if the
farmer faces a particularly bad disease year, the resistant types of
the landrace will survive and yield a harvestable grain. Unfortu-
nately, a clear-cut distinction between landrace and cultivar is
not always maintained. It is not uncommon for a landrace to be
cataloged as a cultivar by one gene bank and as a landrace by
another. While vernacular names are useful for identifying land-
races, they do not have taxonomic standing.
Taxonomy: In Use
As a standardized system for classification of plants, taxonomy
provides a useful way for tracing the origins of grain species. By
knowing the names of wild progenitors and related domesti-
cated species, researchers can make informed decisions when
selecting material for study or for breeding programs. Likewise,
knowing the taxonomic traits by which species are classified
enables researchers to verify the identity of research material.
This is particularly important given the possibility that gene
bank and research collections may hold and distribute misla-
beled or misidentified wild and domesticated germplasm.
The wheats offer a practical example of the importance of
taxonomy in the selection and use of research material.
Because there are several conflicting taxonomic concepts for
the wild and domesticated wheats, a seed request may not be
filled with the taxon actually requested. A general, unspecified
request for accessions of the diploid A-genome progenitor of
tetraploid and hexaploid wheat species may be incorrectly
filled with either T. boeoticum or T. monococcum. While the
wild T. boeoticum and domesticated T. monococcum both have
a form of the A-genome, neither was involved in the evolution
of T. durum (AB genomes) and T. aestivum (ABD genomes)
(Figure 3). The correct species for this request would be
T. urartu. Since T. urartu and T. boeoticum are morphologically
very similar to each other, a researcher, who is unfamiliar
with the taxonomic characters that differentiate them, may
erroneously mistake T. boeoticum for T. urartu.
Taxonomy and Allergies
Taxonomic relationships have a practical application for iden-
tifying potential allergic cross-reactions. For example, a person
with a proven allergic reaction to wheat grain would be more
likely to have similar problems with rye but less with the more
distantly related cereal grain species such as rice or maize
(Table 1). This consideration is particularly relevant to celiac
disease, for which the prime cause is the gluten proteins found
in wheat. Celiacs are also warned to avoid eating the grain of
barley, rye, and triticale, closely related species in the tribe
Triticeae. Although oats, which are classified in tribe Poeae,
are considered to be less harmful for celiacs, some such

individuals find that they must avoid them. On the other hand,
celiacs may eat foods based on other cereal grains, which are
classified in more distantly related tribes (e.g., maize, millet,
rice, and sorghum) and any of the dicot species. Within the
context of taxonomy, the possible cross-reaction with oats but
not with maize, millet, rice, and sorghum is understandable.
The tribes Triticeae and Poeae are located in the same subfam-
ily, the Pooideae. On the other hand, maize, millet, and sor-
ghum are classified in the subfamily Panicoideae and rice in
the subfamily Ehrhartoideae.
Attempts at prediction of allergic cross-reaction may not be
valid if making comparisons with respect to different vegetative
or reproductive parts of the plant and different routes into the
body for example, the inhalation of pollen versus the inges-
tion of grain. It may thus be possible for a person, who is
allergic to wheat pollen, to suffer no adverse reactions to the
wheat grain. However, there is at least anecdotal evidence that
individuals, who experience mild ingestion intolerance to
cereal grains, may have these symptoms exacerbated in pollen
season. Also, allergic cross-reactions are observed between pol-
lens of different species, implying that sensitization by one
species can confer allergy to others. Thus, a positive test reac-
tion to pollen from one species does not necessarily mean that
it is the one causing clinical symptoms, nor need it have been
the one responsible for the original sensitization. The real
culprit may be another cross-reacting pollen. Here, taxonomy
offers a practical tool for elucidating such situations. For the
monocot grains, only the grass family Poaceae is necessary to
consider. Within the very diverse dicot group of pulses, pseu-
docereals, and oilseeds, the higher taxonomic rankings above
Family are of practical use for evaluating allergic cross-
reactions. In providing only a brief introduction into the tax-
onomy of grain species, Table 3 illustrates the hierarchical
relationships that should be considered in dealing with poten-
tial cross-reactions.
Future Trends in Taxonomy
As is evident in the previously mentioned discussion, taxon-
omy is subject to conflicting, and often changing, interpreta-
tions. When taxonomic disagreements prevail, the focus shifts
away from taxonomys real value in aiding communication
among scientists. For example, a long-running debate among
wheat researchers dating from the 1950s over how to classify
wild and domesticated taxa by evolutionary relationships has
created a confusing situation with many different classifica-
tions and no consensus for names or species concepts. This
controversy illustrates the possible problems that lie ahead
should the current trend continue among molecular systema-
tists and research geneticists to restructure traditional taxon-
omy of lower-ranking groups into a natural system that
displays phylogenetic relationships even when they are not
fully resolved and/or open to widely varying interpretations.
Because concepts of plant phylogeny are changing rapidly with
the sophisticated tools of molecular biology, an attempt to
structure taxonomy on molecular evidence may lead to a situ-
ation of constant taxonomic flux. Traditionalists argue that
taxonomy should continue as it now stands guided by the
ICBN. The current approach of the Angiosperm Phylogeny
THE BASICS | Taxonomic Classification of Grain Species 39
Group and its followers offers a viable example of combining
the two systems: structuring higher-order relationships in an
ordinal phylogenetic system while drawing on the stability of
the Linnaean system for treating the lower ranks (Family
GenusSpecies) and revising their classifications as robust
phylogenetic evidence becomes available.
Exercises for Revision
What is the function of scientific names for the grains and
why are they preferred over common names in research,
germplasm conservation, and communication among
scientists?
How do common names convey cultural and historical
information about the societies that cultivated domesti-
cated grains? What role have common names played in
the taxonomic naming system?
What are the distinct differences among these taxonomic
ranks: family, genus, species, subspecies, variety, and culti-
var? In what case is variety and cultivar used interchange-
ably and why is this practice not advisable?
Why is the ICNCP important as a separate set of rules for
naming and classifying domesticated grains? For grain spe-
cies, what types of issues is the ICNCP able to better address
than possible with the ICBN?
How does taxonomic classification help in distinguishing
between cereal and dicot grains and the species classified
within each of these two major grain groups?
With the much-enhanced ability to elucidate phylogenetic
relationships with data from molecular genetic studies,
what types of challenges does the traditional Linnaean
system face when incorporating this information into tax-
onomic classification?
How do competing taxonomic classifications create diffi-
culties for grain researchers? What types of solutions can be
developed to address the controversies associated with tra-
ditional versus natural phylogenetic classifications?
How do evolutionary groupings of grain species in taxo-
nomic classifications aid in assessments of potential allergic
cross-reactions?
Since landraces are often mixtures of multiple species, what
are the challenges faced in cataloging and classifying land-
race diversity? Why is it important to differentiate among
individual types within a single landrace collection?
Exercises for Readers to Explore the Topic Further
Explore the advantages and conflicts posed by the applica-
tion of molecular technologies to taxonomy.
For example, examine and expand on the statement in
Future Trends: molecular systematists (may) suggest
other naming systems and other ways of classifying, which
could lead to a re-invention of taxonomy.
Refer to the article on variety identification, which describes
the use of protein composition and DNA analysis to differ-
entiate cultivars in agriculture. Consider how these
methods might be expanded to compare a wider range of
taxonomic relationships.
40 THE BASICS | Taxonomic Classification of Grain Species
See also: Breeding of Grains: Barley: Genetics and Breeding;
Canola/Rapeseed: Genetics and Breeding; Food Grains:
Intolerance, Allergy and Diseases: Celiac Disease; Genetics of
Grains: Maize: Genetics; Rice: Genetics; Wheat Genetics; The Cereal
Grains: Cereals: Domestication of the Cereal Grains; The Legumes
and Pseudocereals: Amaranth: Overview; Buckwheat: Overview;
Pseudocereals: Overview; The Oilseeds: Oilseeds: Overview.
Further Reading
Angiosperm Phylogeny Group (1998) An ordinal classification for the families of
flowering plants. Annals of Missouri Botanical Garden 85: 531553.
Angiosperm Phylogeny Group II (2003) An update of the Angiosperm Phylogeny Group
classification for the orders and families of flowering plants: APG II. Botanical
Journal of the Linnean Society 141: 399436.
Angiosperm Phylogeny Group III (2009) An update of the Angiosperm Phylogeny Group
classification for the orders and families of flowering plants: APG III. Botanical
Journal of the Linnean Society 161: 105121.
Brickell CD, Alexander C, and David JC, et al. (eds.) (2009) International Code of
Nomenclature for Cultivated Plants, 8th edn. In: Regnum Vegetabile, 8th edn.,
151: Belgium: Drukkerji Geers Scripta Horticulturae 10.
Hanelt P and Institute of Plant Genetics and Crop Plant Research (eds.) (2001)
Mansfelds Encyclopedia of Agricultural and Horticultural Crops (Except
Ornamentals) Germany: Springer-Verlag, English edition.
Heywood VH, Brummitt RK, Culham A, and Seberg O (2007) Flowering Plants of the
World. Ontario, Canada: Firefly Books.
Judd WS, Campbell CS, Kellogg EA, Stevens PF, and Donoghue MJ (2007) Plant
Systematics: A Phylogenetic Approach, 3rd edn. Sunderland, MA: Sinauer Associates.
Linnaeus C (1753) Species Plantarum. Sweden: Laurentius Salvius.
Mangelsdorf PC (1953) Wheat. Scientific American 189: 5059.
McNeill J, Buck WR, and Burdet HM, et al. (eds.) (2012) International Code of Botanical
Nomenclature (Melbourne Code). In: Regnum Vegetabile154: Konigstein, Germany:
Koeltz Scientific Books.
Stuessy TF (2009) Plant Taxonomy: The Systematic Evaluation of Comparative Data,
2nd edn. New York: Columbia University Press.
Vietmeyer ND (ed.) (1989) Lost Crops of the Incas Washington, DC: National Academy
Press.
Watson L and Wrigley CW (1984) Relationships between plants relevant to allergy.
Medical Journal of Australia 141(5 Suppl.): S18.
Wiersema JH and Leon B (2013) World Economic Plants: A Standard Reference, 2nd
edn. Boca Raton, FL: CRC Press.
Williams JT (ed.) (1995) Cereals and Pseudocereals London: Chapman and Hall.
Relevant Websites
http://www.ars-grin.gov The United States Department of Agriculture website for GRIN
(Germplasm Resources Information Network) is set up for taxonomic queries on
holdings of economic plants and with links to other relevant websites for the
taxonomy of economic plants. Economic plant listings include vernacular names
and economic uses. Information on this website is drawn from World Economic
Plants: A Standard Reference.
http://www.eti.uva.nl The Expert Center for Taxonomic Identification (ETI) is an on-
line taxonomic database service supported by an international consortium. The site
hosts a searchable taxonomic database for taxonomic hierarchies, species names,
synonyms, and descriptions.
http://www.iapt-taxon.org/nomen/main.php The online version for the most
recent edition of the ICBN (2012), which is known as the Melbourne Code, is in
English.
http://www.ildis.org Location for the International Legume Database and Information
Service (ILDIS), this website has a searchable database for species information
about the grain pulses.
http://www.ipni.org The International Plant Names Index (IPNI) is a comprehensive
online database of plant names and bibliographic information.
https://www.ksu.edu.wgrc.Taxonomy.taxintro.html This website has comprehensive
listings of historical and current taxonomic treatments of the wheats.
http://www.mobot.org/MOBOT/research/APweb/welcome.html The Angiosperm
Phylogeny Group website has up-to-date revisions to the Angiosperm phylogenetic
trees, extensive reviews of current evidence, and references.
http://www.plantnames.unimelb.edu.au The Multilingual Multiscript Plant Name
Database has a searchable database for locating scientific and vernacular names in
all applicable languages for a particular grain species.
http://tolweb.org.tree.phylogeny.html The Tree of Life Web Project is a searchable
internet source for information on biodiversity, phylogeny, and characteristics of
biological groups.
http://www.umsl.edu.services.kellogg.gpwg.default.htm As a collaborative group of
systematic botanists, the Grass Phylogeny Working Group (GPWG) produced a
phylogeny of the grass family by combining existing data sets. Although GPWG is
no longer active, this website provides access to its results.
http://upov.int/about/en/benefits_upov_system.html The International Union for the
Protection of New Varieties of Plants is an intergovernmental organization that
oversees the convention for protecting intellectual property rights to new plant
cultivars introduced into commerce.
 Grain: Morphology of Internal Structure
H Corke, University of Hong Kong, Hong Kong, Pokfulam, China; Hubei University of Technology, Wuhan, China
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Grain structure determines the necessity for dehulling and
the consequences of wet and dry milling.
The major cereal grains (wheat, rice, and maize), although
botanically similar, have significant differences in organi-
zation of starch and protein reserves.
Wet milling results in separation of major grain compo-
nents; dry milling is generally a size reduction process.
Learning Objectives
To be able to visualize and explain the major anatomical
features of cereal grains.
To understand the distribution of major nutritional com-
ponents in the grains.
To understand how the specific structure of different cereal
grains drives the choice and consequence of wet and dry
milling.
Introduction
This is a wide topic because virtually every aspect of grain
technology is related to grain structure. Handling, processing,
and utilization of grains depend on specific details of their
structure. For example, in dry milling of rice and wheat,
removal of the bran layers is necessary. The wheat grain is not
covered with a tightly attached hull, so unlike in rice, initial
dehulling is not required. Bran removal in wheat is part of the
process of producing fine particulate flour by crushing the
endosperm. Bran removal in rice should leave an unbroken
polished rice grain. In both wheat and rice, endosperm should
be at a specific hardness to achieve the desired outcome. Wet
milling is a fractionation process. In maize, starch, oil, and
protein components are separated. Again, the physical struc-
ture of the grain is exploited in the design of the milling
process. Issues such as these will be covered in detail in this
article. More specifically, the following aspects will be covered:
wheat grain morphology and its effect on dry and wet milling,
rice grain morphology and its effect on dry milling and
nutrition, maize grain morphology and its effect on handling
and wet milling, barley structure and its effect on brewing, and
a comparative summary of general effects of grain structure.
Wheat Grain Morphology and Its Effect on Dry and
Wet Milling
The relationship of the grain to overall spike structure is impor-
tant in milling (Figure 1). Historically, the wheat ancestors
(e.g., the diploid Triticum monococcum) had a brittle rachis
trait. This means that after ripening of the grain and drying of
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00010-3
the plant, the rachis (where the spikelet is attached to the axis)
breaks naturally and the seed (still surrounded by a tightly
bound palea, lemma, and awn) falls from the plant. This is
an essential survival feature for a plant whose seeds are dis-
persed in nature through the action of wind, insects, etc. In a
farmers field, a brittle rachis would not be viable, as harvesting
would be extremely difficult. The genetic step from brittle to
nonbrittle rachis is the key to domestication of small-grain
cereals (e.g., wheat, rice, and barley). However, wheat has a
further useful adaptation to domestication, which differs from
rice. In wheat, the palea and lemma naturally separate from the
grain this is called a naked grain characteristic. In rice, the
grain is covered, that is, the hull is attached to the grain and
requires a separate processing stage (dehulling) to remove it.
Bran
The wheat grain has been described as a single-seeded fruit
called a caryopsis, in which the ripened ovary wall is fused to
the seed. The classical diagram for hexaploid common (bread)
wheat (Triticum aestivum) grain structure was produced by the
Wheat Flour Institute, Washington, DC (Figure 2). The objec-
tive of flour milling is to remove the embryo (germ) outer
layers (epidermis, hypodermis, cross cells, tube cells, seed
coat, and nucellar tissue), which collectively are termed the
bran. The aleurone layer, part of the endosperm, is also
removed with the bran and from a millers point of view may
be included as a bran layer. Much of the vitamin and mineral
content of the grain is contained in the bran. In many coun-
tries, wheat flour is enriched or fortified to replace some of the
nutrients lost in milling. Removal of the bran (which, includ-
ing the aleurone layer, comprises  15% of the grain by
weight) permits production of white flour. Apart from appear-
ance, white flour has different technological properties to
whole-wheat meal (that produced by grinding the whole
grain without bran separation). The presence of large amounts
of bran dilutes the proportion of gluten in the product and
results in a denser, more fiber-rich product. The presence of
small amounts of bran (as bran specks) results in off-white
color in flour and in many final products (e.g., steamed bread).
In fresh white noodles, discoloration to gray is associated with
polyphenol oxidase (PPO) content of flour, and most PPO is
present in bran. In Chinese steamed bread, formation of
brown spots on the surface may also be associated with PPO
effects from bran contamination (and the use of NaCl in the
formulation). Because of the high mineral content of bran
relative to endosperm, bran content in flour can be estimated
by ashing (furnace heating) a flour sample and weighing the
residue. For the same flour yield, a variety that gives lower ash
content (less bran) is of course preferable.
In wheat flour milling, grain is tempered to a uniform
moisture content (usually 15.5%). The effect of this is to
make the bran layers flexible so that under the crushing action
41
42 THE BASICS | Grain: Morphology of Internal Structure

Awn
Spikelet (not always
present)
Rachis
Palea Pales
Lemma Floret
Grain

Rachilla
Glume
(a) (b)

Figure 1 The structure of the wheat spike (ear). Cultivated wheat requires a nonbrittle rachis so that mechanical harvesting is possible and a
loosely attached palea and lemma so that a dehulling step is not required. Reproduced from Evers AD and Bechtel DB (1988) Microscopic structure of
the wheat grain. In: Pomeranz Y (ed.) Wheat: Chemistry and Technology, 3rd edn., vol. 1, pp. 4795. St. Paul, MN: American Association of
Cereal Chemists.

of roller milling, bran will break into large flakes, not be Endosperm
crushed into powder. This formation of flakes of bran results
The product of wheat dry milling is flour, the result of crushing
in pieces that are of different sizes, shapes, and densities to the
endosperm. The endosperm consists of cells surrounded by a
endosperm particles, which on size reduction become flour
pentosan-rich cell wall (mostly arabinoxylans) and contains
particles. Without this physical flexibility in bran, milling
starch granules embedded in a protein matrix. The size of the
would not be possible. However, with crushing, some endo-
cell and the thickness of the cell wall affect the proportion of cell
sperm is likely to be attached to some of the bran flakes.
wall material relative to starch and protein in the endosperm.
Continuing milling to remove the last particles of endosperm
The cell wall absorbs water during dough mixing, so it is signif-
(i.e., aiming for a high flour yield or percentage yield of flour
icant in determining optimum water absorption of flour. The
relative to starting weight of grain) will increase contamination
dimensions of the cell wall are genetically variable. The starch
of flour with bran. The flour yield also depends on several
granules consist of two types, larger A granules and smaller B
other properties of bran:
granules. Starch composition (proportion of amylose to amy-
1. The thickness of the bran layers, which is under genetic lopectin in starch) determines physical properties of any gela-
control tinized starch product resulting from cooking the flour. This is
2. The size of the grain (bran will be a relatively lower fraction most important in Japanese white-salted noodles (udon) where
of a larger grain; also, a plump well-filled grain will be a low-amylose (partial waxy), high-swelling starch gives favor-
preferable) able texture. In many other products, such as baked bread, the
3. The characteristics of the crease starch composition, within normal limits for wheat flour, does
not have much impact on product quality. Starch granules are
The crease in a wheat grain is the lateral fold (Figures 2 and 3)
subject to shear or breakage, termed starch damage in milling, if
ending with the pigment strand. A deep crease results in more
the endosperm is too hard (e.g., genetically hard texture or too
problems with bran removal in milling more bran particles
low moisture content in milling) or if milling equipment is not
attached to endosperm chunks usually result from a deep
adjusted correctly. A broken granule, exposing the interior, is
crease. Also, the crease can trap contaminants, for example,
much more readily subject to hydrolysis by amylases during the
dust and fungal spores, which can cause problems in storage
bread-making process (during fermentation and early stages of
and in cleaning the grain before milling.
baking before the enzyme is inactivated). Some starch damage
Grain color in wheat, usually due to red pigmentation in the
may be desirable to provide a substrate for yeast growth, but
bran (seed coat), is usually classified as red (the presence of
excessive damage adversely impacts on quality and may result in
pigmentation) or white (absence). White wheat flour, if contam-
a sticky texture in the final loaf. Endosperm hardness is largely
inated with bran, will show less discoloration than red wheat
due to the structure of two puroindolines (components of fria-
flour. This is why hard white-type wheat is becoming more
bilin), which are amyloplast membrane proteins present in the
widely accepted as suitable for Asian noodles. Other colors are
endosperm. There is a widespread confusion about grain hard-
possible but rare in commercial trade for example, blue, pur-
ness (protein-influenced) and noodle texture (starch-
ple, or even black wheat, with a dark purple pigment in the seed
influenced). A high-swelling starch, but not necessarily a soft
coat, which is used in some specialty products in China.
 THE BASICS | Grain: Morphology of Internal Structure 43

Hairs of
brush

Endosperm
Cell filled with
starch granules
in protein
matrix
Cellulose walls
of cells
Aleurone cell
layer (part of
endosperm but
separated
with bran)
Nucellar tissue
Seedcoat
(testa)
Tube cells
Cross cells
Hypodermis
Epidermis

Scutellum

Sheath of shoot

Rudimentary
shoot
Rudimentary
primary root
Root sheath
Root cap

Longitudinal section of grain of wheat

(enlarged ~35 times)

Crease
Endosperm

Pigment
strand
Bran
Germ

Cross-sectional view
Figure 2 The structure of the wheat grain longitudinal and cross-sectional views. Reproduced from Pomeranz Y (1987) Modern Cereal Science
and Technology, pp. 2539. Weinheim, Germany: VCH Publishers.
44 THE BASICS | Grain: Morphology of Internal Structure

Outer kernel layers Central starchy endosperm Crease region

Wheat 100 m 100 m 100 m

Barley 100 m 100 m 100 m

Oat 100 m 100 m 100 m

Rye 100 m 100 m 100 m

Figure 3 Outer kernel layers, central starchy endosperm, and crease region of kernel cross sections of wheat, barley, oat, and rye stained with acid
fuchsin calcofluor. Reproduced from Dornez E, Holopainen U, Cuyvers S, et al. (2011) Study of grain cell wall structures by microscopic
analysis with four different staining techniques. Journal of Cereal Science 54: 363373.

endosperm, is required to make a soft-textured noodle (such as
Japanese udon).
Embryo (Germ)
The embryo consists of the rudimentary structures (shoot and
root) of the plant, which can grow from the grain, plus the root
and shoot sheathes, the root cap, and the scutellar tissue. The
embryo, 3% of the grain weight, contains  8% of the grain
protein, but it is mostly in the form of enzymes related to
subsequent growth of the seedling, not storage protein (gluten)
as in the endosperm. Thus, the wheat germ protein is of higher
quality (more favorable amino acid balance, such as higher
lysine content) for human nutrition. The lysine level in protein
in the embryo is 8%; in endosperm protein 2%, and in
bran protein 4.5%. In addition, 20% of the lipids of the grain
are in the embryo (oil-containing spherosomes are present in
the aleurone layer and in scutellar cells), but there are also high
levels of lipase and lipoxygenase enzymes, which hydrolyze
and oxidize oil, causing rancidity. Wheat germ hence has uses
as a food material or supplement for human nutrition and is
valuable as an additive to animal feed (particularly for

monogastric animals such as pigs and poultry). On the other
hand, wheat germ, unless treated, has a short shelf life. Whole-
wheat meal also has a shorter shelf life than white flour.
Removal of germ to produce flour results in a product that is
stable for long periods of time.
Other Features
Dry milling of durum (Triticum durum) wheat (a tetraploid
species) to make semolina (larger particle size than flour
from common wheat) has a few special features related to the
durum endosperm texture. The physical hardness (vitreous-
ness) of durum endosperm makes the grain liable to break
during combine harvesting. These broken grains should be
separated and recovered during cleaning of grain before mill-
ing, or a high level of wastage may occur. Optimum tempering
of durum grain before milling is also essential and varies with
the vitreousness of the endosperm. Careful milling (e.g., more
break rolls than for bread wheat) is needed to achieve suitably
fine semolina for high-quality pasta.
In wheat wet milling (to produce wheat starch, gluten, and
wheat germ oil), separation is by soaking the grain, crushing,
and sedimenting off the starch component. B-type starch gran-
ules are too small to easily sediment, and their complete
removal would require expensive centrifugation steps. Hence,
Awn
Lemma
Palea
Hull
Sterile
lemmae
Rachilla
Figure 4 The structure of the rice grain. Reproduced from Juliano JB and Aldama MJ (1937) Morphology of Oryza sativa L. Philippines
Agriculturalist 26: 1134; Juliano BO (1993).
THE BASICS | Grain: Morphology of Internal Structure 45
a genetically low proportion of B-type compared to A-type
granules would be a strong advantage for increased starch
yield in wet milling. Considerable effort is being expended to
identify more suitable wheat varieties for this use.
A significant recent development has been the application
of Raman microscopy (in combination with further develop-
ment of selective staining for optical microscopy) to image and
localize the presence of, for example, b-glucans, protein, starch,
and ferulic acid in wheat and barley grain. The advantage of
Raman microscopy, applied alone, is the absence of required
pretreatment or staining. An example of the capability of high-
quality selective staining can be seen in Figure 3; such infor-
mation as the cell wall size and thickness, the size and packing
of starch granules, the nature of the bran layers, and the geom-
etry of the crease can be invaluable for genetic improvement of
wheat.
Rice Grain Morphology and Its Effect on Dry Milling
and Nutrition
Grain Structure
The anatomical structure of rice (Figure 4) is of course very
similar to that of wheat. However, grain shape is of more
significance. Short-, medium-, and long-grain rice are common
Pericarp
Seedcoat
Nucellus
Aleurone layer
Subaleurone layer
Endosperm
Starchy endosperm
Scutellum
Plumule
Embryo
Radicle
Epiblast
46 THE BASICS | Grain: Morphology of Internal Structure
classifications. In the United States, texture can be recognized
from grain length short grain is low amylose (sticky) and
long grain is intermediate amylose (2025%) (harder cooked
texture such that grains are separate after boiling). This is a
marketing phenomenon, not a genetic relationship, and in
other countries, long grain may well be waxy or sticky types,
sometimes a source of confusion to traveling Americans. Grain
shape (expressed as ratio of length to width) affects milling. A
rounded grain with a low length/width ratio will be more
resistant to breakage than a slender type of grain. Rice lacks
the crease found in wheat grains. This enables abrasive milling
to remove the bran and leave only the endosperm, greatly
simplifying the milling process.
The hull, which covers the rough (also known as paddy)
rice, comprises 1724% of the weight of the paddy rice. It
consists of lemma and palea, awn, the rachilla, and lemmas.
Varietal differences in percentage of hull affect yield of brown
rice, and this in turn affects milled rice recovery (yield of white
polished rice). The hull (husk) is silica-rich and indigestible for
human nutrition, even after heat treatment, even though it is
rich in minerals.
Cracking
In producing polished white rice, the control of cracking is
essential. Cracks caused by poor drying conditions after har-
vest, for example, will result in grain that breaks easily in
milling (dehulling or polishing) producing a lower yield of
head rice and consequently much lower value product. In the
production of parboiled rice, rice with the hull still attached is
100
80
60
(%)
40
Fiber
Ash
20 Fat
20
Outer layer
Proportion of the kernel
Figure 5 The effect of progressive surface abrasion of brown rice on content of major constituents. Reproduced from Barber S (1972) Milled rice and
changes during aging. In: Houston DF (ed.) Rice: Chemistry and Technology, pp. 215263. St. Paul, MN: American Association of Cereal Chemists;
Juliano BO (1993).
steamed, driving nutrients from the hull and the bran into the
endosperm, inactivating enzymes that would cause rancidity,
and reducing microbial load. However, after steaming, the
parboiled rice must be dried slowly before it is dehulled and
polished, or cracks will form with the same effect as before.
Parboiled rice, despite the steaming treatment, does not
require less time to cook by the consumer than ordinary
white rice, because water penetration to the center of the
grain is slow. Instant rice is prepared by steaming polished
white rice. It is dried rapidly, forming numerous internal
cracks. These cracks, not the starch gelatinization induced by
steaming, allow the product to be cooked by the consumer in a
few minutes because they aid water penetration. Instant rice is
more physically liable to breakage in transportation than raw
polished white rice, and the softer texture resulting from the
cracks is unattractive to traditional rice consumers in many
parts of the world.
Rice Bran Layers
The bran layers (pericarp, seed coat, nucellus, and aleurone
layer) contain most of the minerals (represented as ash), vita-
mins, fiber, fat, and much of the protein present in the rice
grain (Figure 5). Considerable debate over the years has taken
place over the suitability of white polished rice, compared to
brown rice, in nutritionally deficient diets in some developing
nations. In fact, bran also contains phytate (phytic acid), which
binds to available minerals provided by brown rice, and may
also bind and remove additional minerals from the gastroin-
testinal tract. Brown rice consumption may therefore cause a
Starch
Protein
40 60 80 100
Center

Compound
starch
granule
Figure 6 The rice endosperm indicating structure of spherical and crystalline protein bodies. Reproduced from Coffman WR and Juliano BO (1987)
Rice. In: Olson RA (ed.) Nutritional Quality of Cereal Grains: Genetic and Agronomic Improvement. Madison, WI: Agronomy Society of America;
Juliano BO (1993).
net loss of minerals to the body and in addition is susceptible
to rancidity in storage (lipids and lipases still present) and
more susceptible than polished rice to storage pest attack.
Rice Endosperm
Rice starch granules are small (24 mm), polygonal, and com-
pound, that is, many small granules are compacted together.
Compositionally, rice starch varies from 0% (waxy, sticky, and
glutinous) amylose to over 35%. This range is much wider than
found naturally in hexaploid wheat starch. Expression of natu-
rally occurring waxy mutants is more likely in a diploid cereal
(e.g., rice, maize, and sorghum) than in a hexaploid where the
effect is buffered by the other two sets of chromosomes present.
Rice differs from wheat and maize in having crystalline protein
bodies rich in glutelin, in addition to the prolamin-rich protein
bodies common to all cereal endosperms (Figure 6). The aleu-
rone layer (considered here as the botanical part of the endo-
sperm, although it is processed as part of the bran) differs in
thickness (usually one to seven cell layers) depending on vari-
ety. The aleurone cells contain protein bodies and lipid bodies.
Rice Embryo
The rice embryo is smaller relative to the whole grain than in
wheat. It contains several types of parenchymatous cells, con-
taining protein bodies and/or lipid bodies. The main parts of
the embryo are the cotyledon and the embryonic axis. The
plumule and radicle are similar to other cereal grains. The
high lipid content of the embryo contributes to susceptibility
to rancidity in storage of brown rice.
Maize Grain Morphology and Its Effect on Handling
and Wet Milling
Maize is monoecious, having separate male and female flower-
ing parts on the same plant. The female seed-bearing part of the
plant (called the ear) develops off the lower-middle side of the
THE BASICS | Grain: Morphology of Internal Structure 47
Large
spherical
Crystalline Protein
bodies
Small
spherical
1 m
plant and consists of a central cob with 800 seeds attached in
rows by the pedicel and covered by a husk. The pedicel acts as a
conduit for photosynthetic products supplying the developing
grain. The male part of the flower the tassel produces pollen
and is at the top of the plant. The maize grain (Figure 7) is 10
times larger than small-grain cereals, an average dent maize
grain weighing 300 mg. The flattened sides of the mature
grain are due to pressure from the tight packing in rows on
the cob. After removal of the husk, the maize grain is exposed
(naked characteristic) and easily separated from the cob.
Handling
Shelled maize grain (grain separated from the cob and packed
in bulk) is more likely than other grains to enter export mar-
kets. A high proportion of the US maize harvest is exported,
often to highly quality-conscious markets such as Japan. A
typical export grain may enter the transportation system by
traveling by barge down the Mississippi, entering the Gulf of
Mexico for the long voyage to Asian markets. This is physically
stressful on the grain, and a poorly dried grain with internal
cracks is liable to breakage. This contributes to quality loss and
increased susceptibility to insect or fungal attack and moisture
damage. The primary criteria for successful shipment of maize
are suitable endosperm texture (hard enough) and suitably
controlled harvesting and drying to avoid cracking.
Wet Milling
Most maize that is not fed to animals is processed by wet
milling to produce starch (with many diverse applications,
such as further processing high fructose corn syrup, bioetha-
nol, or modified starch for food and nonfood industries), oil
(a high-quality cooking oil), and protein or fiber-rich fractions
that are used in livestock feed. The composition of these com-
ponents in the grain is of course of the utmost importance. As
the maize seed is much larger than wheat or rice, the relative
proportion by weight taken up by the bran layers (including
aleurone layer) (5%) is less than in these other cereals.
48 THE BASICS | Grain: Morphology of Internal Structure

Hull

Epidermis
Mesocarp
Cross cells
Tube cells
Seedcoat (testa)
Aleurone layer
(part of endosperm
but separated with bran)

Horny endosperm

Floury endosperm

Cells filled with starch

granules in protein matrix

Walls of cells

Scutellum
Plumule or rudimentary
shoot and leaves
Longitudinal section
of a grain of corn Radicle or
(enlarged -30 times) primary root

Tip cap

Scutellum Embryonic axis

Pericarp

Horny Floury
endosperm endosperm Cross-sectional view
Figure 7 The structure of the maize grain longitudinal and cross-sectional views. Reproduced from Pomeranz Y (1987) Modern Cereal Science
and Technology, pp. 2539. Weinheim, Germany: VCH Publishers.

The embryo of maize is a much higher proportion of the
grain (11%, containing 33% of the oil) than in other
cereals; hence, the overall oil content of the grain is quite
high (typically 5%). Also, a higher proportion of the total
protein is nonstorage in nature (e.g., enzymes of the embryo),
so overall, the protein of maize is higher quality for human
nutrition than that of wheat flour. However, maize protein,
like other cereals, is deficient in lysine. High-lysine mutants
(opaque-2) give more favorable amino acid balance suitable
for feeding people or monogastric animals even without
additional protein sources. In practice, spread of opaque-2 vari-
eties has been limited because in most cases, production of
normal endosperm maize, and supplementation with soy pro-
tein, is easier for animal feed production. Similarly, a classic
long-term selection experiment at the University of Illinois
showed that the oil content of maize could be increased to
extremes of over 40%. This was basically a result of increased
embryo size relative to endosperm.
The maize endosperm, the source of starch and most of
the protein, has several interesting features. Maize types are

classified by endosperm hardness. Floury is a soft endosperm
that can be easily crushed to a friable powder. It is physically
susceptible to storage pest (mainly insect) and fungal attack.
An example of floury maize is the opaque-2 mutant. Dent has a
high proportion of floury endosperm, but some horny or hard
endosperm (Figure 7) and is the most widely produced com-
mercial maize. The dent refers to the indentation at the top of
the grain on drying when it contracts due to the floury endo-
sperm component. Flint maize (of which popcorn is one
special type) has only horny and no floury endosperm. The
top of the seed is rounded, not indented, and it is physically
very hard, even after cooking meal prepared from it.
The outer layers of maize (the seed coat in particular)
contribute the color, which is white or yellow in nearly all
commercial maize production. Mass production of maize for
feed and industrial use, for example, in the United States, is
predominantly yellow dent. Some white dent is produced for
breakfast cereal (e.g., corn flake) production (which follows
maize dry milling). The carotenoids (xanthophylls) of yellow
maize are nutritionally important and also, in poultry feed,
contribute yellow color to egg yolks and to skin and fat of meat
poultry. People in regions where maize is a daily food staple
may have strong color preferences; for example, in southern
Africa, white (dent) maize is strongly preferred because yellow
maize is historically associated with flintiness (although there
is no genetic relationship).
Maize Dry Milling
This also requires a physically sound clean sample with appro-
priate hardness. Dry milling in the United States usually
implies use of white maize (a minor proportion of total US
production) to make endosperm chunks, which may be
steamed and flaked to make corn flakes. Larger dry milling
industries exist to serve the needs of maize tortilla production
(Mexico) and maize porridge production (a staple diet in
much of sub-Saharan Africa) requiring maize meal.
Barley Grain Structure and Effect on Malting
The barley grain is similar in structure to the wheat grain. For a
diagram and further information on this topic, see elsewhere in
this encyclopedia.
Overview
The same principles apply when considering the structure
and utilization of other grains, such as sorghum, millets,
oats, or even the pseudocereals buckwheat and Amaranthus.
First, consideration should be the required product, flour,
separation of starch/protein/oil, or malt, which drives the
choice of process wet or dry milling or malting. The size
and shape of the grain, particularly the dimensions of the
outer layers will mostly affect dry milling. In some cases, the
seed is extremely small (e.g., with Amaranthus) and dry mill-
ing flour yield will be low. For such crops, selection for larger
grain size can greatly increase flour yield. For wet milling, the
fraction of most value (usually starch) will drive the design
of the separation process. Usually, the starchprotein
THE BASICS | Grain: Morphology of Internal Structure 49
binding needs to be loosened by soaking in a solution,
such as in alkaline wet milling, where the physical properties
of the starch may be affected. Care should be taken that the
fractions separated still achieve proper functionality in the
intended use. For a useful further overview of the develop-
mental aspects of cereal grain structure, the reader may refer
to Evers and Millar.
Exercises for Revision
Draw a typical cereal grain representing the major anatom-
ical features common to all cereal species.
Explain how starch, protein, and oil are stored in different
cereal grain species.
Exercises for Readers to Explore the Topic Further
Investigate the concept of domestication for a selected
cereal grain crop, showing the necessary and desirable
changes from wild relatives to modern cultivated plants.
Design an ideal maize grain for different processing appli-
cations (e.g., bioethanol and high oil), and indicate what
genetic approaches have been and could be taken to pro-
duce it.
The bran is often discarded after processing. What unusual
and interesting variants in bran compositional features exist
(e.g., in pigmentation), and how could these be better
exploited for value-added uses?
See also: Barley, Rice and Maize Processing: Barley: Malting;
Maize: Wet Milling; Beverages from Grains: Beverages: Distilled;
Breeding of Grains: Maize: Breeding; Food Grains and the
Consumer: Fortification of Grain-Based Foods; Genetics of Grains:
Maize: Genetics; The Basics: Grain and Plant Morphology of Cereals
and how characters can be used to identify varieties; The Cereal
Grains: Rice: Overview.
Further Reading
Barber S (1972) Milled rice and changes during aging. In: Houston DF (ed.) Rice:
Chemistry and Technology, pp. 215263. St. Paul, MN: American Association of
Cereal Chemists.
Coffman WR and Juliano BO (1987) Rice. In: Olson RA (ed.) Nutritional Quality of
Cereal Grains: Genetic and Agronomic Improvement. Madison, WI: Agronomy
Society of America.
Dornez E, Holopainen U, Cuyvers S, et al. (2011) Study of grain cell wall structures by
microscopic analysis with four different staining techniques. Journal of Cereal
Science 54: 363373.
Evers AD and Bechtel DB (1988) Microscopic structure of the wheat grain.
In: Pomeranz Y (ed.) Wheat: Chemistry and Technology, vol. 1, pp. 4795. St. Paul,
MN: American Association of Cereal Chemists 3rd edn.
Evers T and Millar S (2002) Cereal grain structure and development: Some implications
for quality. Journal of Cereal Science 36: 261284.
Hoseney RC (1994) Cereal Science and Technology, pp. 128. St. Paul, MN: American
Association of Cereal Chemists 2nd edn.
50 THE BASICS | Grain: Morphology of Internal Structure
Jaaskelainen AS, Holopainen-Mantila U, Tamminen T, and Vuorinen T (2013)
Endosperm and aleurone cell structure in barley and wheat as studied by optical and
Raman microscopy. Journal of Cereal Science 57: 543550.
Juliano JB and Aldama MJ (1937) Morphology of Oryza sativa L. Philippines
Agriculturalist 26: 1134.
Juliano BO and Bechtel DB (1985) The rice grain and its gross composition.
In: Juliano BO (ed.) Rice: Chemistry and Technology, pp. 1757. St. Paul, MN:
American Association of Cereal Chemists 2nd edn.
Pomeranz Y (1987) Modern Cereal Science and Technology, pp. 2539. Weinheim,
Germany: VCH Publishers.
Watson SA (2003) Structure and composition. In: Watson SA and Ramstad PE (eds.)
Corn: Chemistry and Technology, pp. 5382. St. Paul, MN: American Association
of Cereal Chemists.
Relevant Websites
http://www.cerealsdb.uk.net/cerealgenomics/WheatBP/Documents/DOC_Milling.php
Website from School of Biological Sciences, Bristol University, with interesting
images information related to wheat processing.
http://www.namamillers.org/education/ Website of the North American Millers
Association has useful information on the relationship between grain structure and
milling for various cereal grains.
http://www.qaafi.uq.edu.au/rice-grain-storage-protein-composition A presentation by
Dr. Daniel Waters of Southern Cross University on aspects of rice grain and protein
structure and composition.
 Grain and Plant Morphology of Cereals and how characters can be used
to identify varieties
J Wyatt, NIAB, Cambridge, UK
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
An introduction summarizing the definition of plant mor-
phology, how it has evolved, and how it can be used to
identify cereal varieties
A table giving a list, in order of importance, of the species
covered in the manuscript
A general section outlining the characters that are shared by
the different cereal species
Specific sections for each species on general growth habit
and inflorescence
How specific parts of a mature grain can be used to identify
many characters
Drawings and photographs to illustrate the text
Identifies some of the differences between the Triticum
species
Learning Objectives
The article can only give a brief introduction to cereal plant
and grain morphology and how the major characters can be
used in the varietal identification of cereals. It is hoped that
this article will lay a foundation for encouraging the student
to partake in further studying cereal plant and grain mor-
phology and how that knowledge gained can be used to
research the many other factors that influence the growing
of cereals across the world.
Introduction
The definition of plant morphology is the physical appearance
of a plant. It can be applied to any species and involves a
detailed study of vegetative and reproductive characters in
order to form a profile of a plant, which can be used to make
general comparisons of plant species displaying a similar
structure or detailed comparisons within a species to identify
varieties (cultivars). Identifying varieties and species of plants
or grains by morphological means is still the main method
used in cereal breeding and varietal identification and provides
the basis for the definitive description of a variety.
Plants within a species share a similar genetic base, and thus,
it is possible to group them into a taxonomic family. Cereals is a
generic term used to encompass the major grain-producing spe-
cies whose products are some of the worlds most important
staple foods and have been since man started growing them for
food some 12 000 years ago. A cereal variety is a population of
plants that can be identified as being clearly distinguishable from
another population by stable inherited morphological charac-
ters. Wheat (Triticum aestivum), rice (Oryza sativa), and maize
(Zea mays) account for over half of the cereals consumed world-
wide. Cereals sit in the family Poaceae (synonymous with
Gramineae). While cereals differ in detail in the way the
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00009-7
characters of each species are expressed, they share a general
commonality in growth habit and flowering parts. By under-
standing the morphology of each species, it is possible to use it
to identify the varieties (cultivars) that exist in each classification.
Recognition of the importance attached to identifying varieties is
highlighted by the existence of UPOV (The International Union
for the Protection of New Plant Varieties). The test guidelines put
together by the 71 members of UPOV provide useful informa-
tion of the major characters used in establishing the morpholog-
ical identification of the different varieties of cereal species.
The following section begins with a description of the main
morphological characters common to all the cereal species and
is followed by further sections specific to each species. Illustra-
tions and photographs are used to help in the identification of
the characters described.
General Cereal Morphology
Cereals are annuals within the family Poaceae (synonymous
with Gramineae). The method of production of the cereal species
is dependent on the climatic/soil conditions, whether the crops
are planted as seeds or as small plants, and whether the variety
requires a cold period for initiation of the flowering process
(vernalization) and subsequent development of the grain.
Cereals are classified as monocotyledons (one embryo leaf
per seed) and share the following characters: an adventitious
root system where numerous roots develop below the surface
from the stem, the early development of stems from the lower
leaf sheaths, and a series of alternating leaves on the stem that
are linear in shape and display parallel veins within the leaf
blade. The flower structure is arranged in threes or multiples of
three (trimerous). The distribution variation, that is, length,
width, curvature, and the frequency of different morphological
characters, is used to distinguish between different varieties.
The morphology of rice and maize shows some variation
from other cereals, having evolved over the centuries by adapt-
ing to suit the climate and locality in which they grow. Exam-
ples of this are the hollow stems of rice that can store air and
the brace roots of maize that act as additional anchorage. The
typical growth habit of (a) maize and (b) other cereals is
shown in Figure 1.
Once the grain has germinated below ground, the sheath
(coleoptile) protecting the first leaf emerges above the soil
surface. As the plant grows, the leaves develop from a growing
point, which remains below the soil surface, with new leaves
emerging from each subsequent leaf sheath. By the fifth to sixth
leaf stage, side shoots (tillers) will start to emerge from the base
of the first leaf (axil). These tillers will go on to form new leaves
in the same manner as those of the first leaf (Figure 2).
The leaves on the primary and secondary tillers grow in a
tubelike structure and break out as a leaf blade at the node. It is
at this juncture that the auricles are found (Figure 3). If
51
52 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties

(n)

(a) adventitious root system

(b) basal leaf sheath if present (m)
(c) leaf sheath
(j) (d i) early leaf formation
(d ii) twisted leaf of barley (l)
(e) tiller emerging from leaf sheath
(a) adventitious roots
(f) node
(b) additional brace roots (k)
(f) (g) auricles if present
(c) soil surface (i) (h) ligules if present
(d) secondary tiller (i) leaf blade
(e) leaf blade margins either (j) stem
straight or lobed (k) flag leaf
(f) flag leaf (l) neck
(e)
(m) ear or panicle (j)
(g) leaf node auricles absent
(h) silk (i)
(i) leaves which cover (h)
emerging cob (g)
(j) tassle

(h) (g)

(e) (f)
(e)

(e)

(d) (d i)
(d ii) (c)
(b)
(c)
(b)

(a)
(b) (a)
(a)

Figure 1 (a) Typical growth habit of maize. (b) Typical growth habit of a cereal.

(a) auricle if present

(b) ligule if present
(c) node
(d) stem/tiller

(d)

(c)

(b)
(d)

(a)
(d)

(d)

Figure 2 Early stem (tiller) formation.

 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 53

hairs present
rice (numerous)
wheat (sparse)
triticale (occasional)

hairs absent, e.g., (b)

barley no auricles
oats
maize
triticum turgidum
secale x secale

(a)
(c)

Figure 3 Auricles and the difference between cereal species.

Weak Strong
(a)

Weak Strong
(b)

Photo 1 (a) Flag leaf glaucosity in winter wheat. (b) Ear glaucosity in winter wheat.
54 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties

present, they encircle the stem and are thought to provide a
barrier within the node to prevent moisture and fungal patho-
gens from entering. Generally, the stems develop as a hollow
stem and continue elongating until the uppermost leaf appears
(flag leaf), from which the inflorescence emerges.
The number of leaves on the stem is dependent on the
eventual height of the plant with the ultimate leaf (flag leaf)
usually smaller than the other leaves. The exterior surface of a
leaf is covered by the cuticle, a structure that helps prevent
moisture loss and the penetration of the external surface by
dirt and harmful microorganisms. Those plants that display a
high level of gray coloration have an extra layer of a waxlike
material (epicuticular wax) on the surface, giving the plant an
additional defense mechanism against moisture loss.
Color differences due to a difference in glaucosity can be
observed in different species and in different parts of the plant,
ranging from green to blue gray in the leaves and stem
(Photo 1(a) and 1(b)). It is also possible to observe different
levels of glaucosity on the glumes and grain. Anthocyanin
coloration (pigment), if present, is found in different levels of
intensity and at different locations within the plant, depending
on species (Photo 2(a)2(c)).

(a)
(b)

Pigment absent Pigment present

(c)

Photo 2 (a) Pigment on grain of barley. (b) Pigment on anthers of maize. (c) Basal leaf sheath barley.
 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
(h)
(g)
(d)
(a)
(e)
(a) glumes
(b) lemma
(c) rachis
(d) ovary
(e) lodicule
(f) stigmas
(g) filament
(h) anther
Figure 4 Inflorescence of a cereal floret.
Each species shares common ground in having an inflo-
rescence made up of a series of florets attached to a central
stem, which contain (in all species) the male and female
reproductive organs (Figure 4). Wheat, rye, triticale, durum
wheat, and barley have a central stem (rachis) with a series
of spikelets attached to it. Each spikelet is made up of
several florets, and once fertilization has taken place, these
florets develop into individual grains arranged alternately
on the stem (Figure 5). The first rachis segment attaches the
ear to the stem. Rice, maize, and oats share a similar inflo-
rescence with a series of branches forming from the upper-
most node (panicle). Panicles can be one-sided (unilateral),
spreading, or triangular (equilateral) in appearance
(Figure 6) with a series of florets lying along the branches.
Maize has a separate male and female inflorescence, where
numerous male florets (tassel) are to be found at the apex
of the plant and the female florets (silk) located within the
stem node(s) (Photo 3).
All cereal species are able to self-pollinate with male and
female flower parts on the same plant (monoecious), although
pollen is accepted from other sources if their own pollen
supply is scarce. Once fertilization has taken place, the front
(ventral view) of the developing caryopsis is protected by the
palea, the back (dorsal view) by the lemma, which overlaps
the palea at the side. In wheat and oats, the whole structure is
protected further by the glumes. In order to assist pollination,
two swollen structures (lodicules), on either side of the ovary,
prise open the lemma and palea. Once fertilization has
taken place, the lodicules shrivel but remain attached to the
lemma base.
For species other than maize, the developing grain may
have a bristle of varying lengths extending from the lemma.
55
(f) (b)
(c)
The structures are known as awns (barley) or beards (wheat,
durum, rye, and triticale) if they are present on each spikelet.
Where they appear in a reduced form on the apex or upper part
of a wheat ear, they are referred to as scurs. Once fertilization
has taken place and providing that no outside influence affects
the structure, the grains will go on to produce viable seeds.
Each caryopsis (grain) is made up of (i) the embryo that
contains all the structures essential for development and (ii)
the endosperm that provides nutrients essential for the germi-
nating seed, mostly in the form of starch. The whole is held in a
covering (pericarp) and remains dormant until the right con-
ditions for germination are present, normally moisture and the
optimum temperature.
Species Specific Plant and Grain Morphology
Rice (Oryza sativa) (Table 1(a))
In terms of human consumption, rice is the most important
cereal grown. There are two subspecies: O. japonica that pro-
duces a short-grained sticky rice and is more tolerant of tem-
perate climates and O. indica that produces a longer-grained,
nonsticky type of rice that has a high moisture and temperature
requirement and is more suited to tropical conditions. There
are two main types: rice that grows on flooded land (paddy)
and deepwater rice that requires more than 50 cm of rain for at
least 1 month (Photo 4). Deepwater rice varieties are capable
of floating in depths from 80 to 100 cm and, by having longer
leaves than the traditional tall paddy varieties, ensure that a
supply of oxygen is available to sustain the whole plant. As the
stems elongate, they are capable of producing their own roots
(nodal roots) and becoming independent from the primary
56 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties

(c)

(a) first rachis segment

(b) rachis segment varies in length
and curvature
(c) sterile spikelet
(d) glumes
(e) collar (d)

(b)

(a) Single line of rachis segments of maize

(b) cob, showing truncation of the rachis

Note: rachis segment of wheat, (e)

durum, triticale, and rye follow
similar format

(a)

Figure 5 (a) Rachis segments of barley. (b) Single line of rachis segments of maize cob, showing truncation of the rachis.

one-sided panicle
(unilateral)
ear
e.g., oat
e.g., wheat
barley many-branched spreading panicle
e.g., rice

Figure 6 Different forms of inflorescence.

 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 57

stem. The stems are especially adapted to contain a large num-
ber of air cells that allow oxygen to pass through to the root
system. A typical rice plant has a hollow stem and will attain an
average height of 11.8 m. Depending on environmental
conditions, the fertility of the soil, and the variety, it can
grow to 5 m in length.
Rice is the only cereal species that is germinated before
planting and set out once the seedling has a minimum of
three leaves. The adventitious roots develop from underground
stem nodes and form the basis of the root system. Whether the
root or the shoot emerges first is determined on whether germi-
nation takes place in aerobic conditions (when the root will
appear first) or in flooded conditions (anaerobic) when the
shoot will appear first. As the main (primary) tiller develops
and if growing conditions are favorable, a secondary tiller will
grow from the primary tiller, followed by a third (tertiary) tiller.
The earlier a variety matures, the shorter the plant and the
distance between internodes. A typical flag leaf encloses the
emerging panicle. The flower (floret) structure differs from
other cereals in having six well-developed stamens with short
filaments but follows other cereals in having a pistil with a
feathery stigma and lodicules (Figure 7). Depending on the
variety, the color of the inflorescence and the subsequent grain
can be gold, brown, light to dark red, purple, or black. Virtually,
all varieties show basal leaf sheath coloration and colored
auricles, which can extend into the lemma tip and keel.

Photo 3 Maize plant. Photo 4 Rice crop.

Table 1 List of the cereal species covered by this article in order of their importance worldwide. The common name is given first, followed by the
Latin declination and a further column, which highlights additional information on the usage of the species. It will be noted that in terms of human
consumption, rice is the most important cereal but that maize is the most important in terms of worldwide production

Species (common
name) Latin name Usage

(1(a)) Rice Oryza sativa (subsp. O. indica and Most important cereal in terms of human consumption
O. japonica)
(1(b)) Maize Zea mays Most important in terms of worldwide production but not all the grain produced goes for
human consumption
(1(c)) Wheat Triticum aestivum Second to rice in terms of human consumption. Both winter and spring types exist
(1(d)) Durum Triticum durum Second most important Triticum species after wheat
(1(e)) Triticale Triticale (X Triticosecale) Hybrid of wheat () and rye () has the yield of wheat with the additional advantage of
tolerance to diseases and environmental changes
(1(f)) Rye Secale X cereale Suited to much poorer soils than wheat and durum
(1(g)) Barley Hordeum vulgare Ranks fourth in importance and used primarily as a food source and as a raw material in
the brewing/distilling industry
(1(h)) Oats Avena sativa A higher tolerance to cool conditions and rainfall
58 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
(f)
(g)
(d i)
(e)
(c)
(d)
(a) pedicel
(b) glumes
(c) palea
(d) lemma (b)
(d i) lemma nerves
(e) feathery stigma
(f) six anthers
(a)
Figure 7 Typical floret of rice.
Maize (Zea mays) (Table 1(b))
In terms of the volume grown, maize is the most important
cereal species, but much is used for purposes other than food,
which places it second to rice in terms of worldwide staple food
consumption. It displays major differences when compared
with the other species covered in this article. The main stem
grows to 1.23 m in height. In the early stages of development,
a mass of underground adventitious roots are produced just
below the soil surface, and as the plant grows, a series of
additional brace roots (Photo 5) emerge above the surface
from the two lowermost leaf nodes. Although the brace roots
have the facility to grow quickly and provide stability for the
plant, maize is still very susceptible to wind damage. Depend-
ing on the eventual plant height, an average of twenty alternate
leaves will develop up the stem.
Maize is classified as a hermaphrodite plant in having the
male and female reproductive organs sited in different loca-
tions on the same plant. The staminate (male) inflorescence
(tassel) develops at the apex of the plant, normally after the
emergence of the 14th or 15th leaf, as a panicle producing a
series of spikelets in which fertile and sterile florets sit. The
pistillate (female) inflorescence (silk) lies normally in the leaf
axils of the 11th13th leaves (Photo 6(a) and 6(b)) with pairs
of spikelets sitting on a much truncated rachis. From each silk,
a series of tubelike structures emerge with small leaves, which
will form the covering of the cob at maturity (husk leaves). At
one end is the undeveloped grain and at the other a feathery
Photo 5 Brace roots of maize.
style. Pollen is released from the male anthers in the tassel and
dispersed by the wind (anemophilous dispersal). As a result of
the relative heaviness of the pollen, it drops the short distance
to the silk below or onto a neighboring plant. Once fertiliza-
tion has taken place, each individual maize grain develops on
the shortened rachis, the whole structure forming the cob
(Photo 7). The color of the cob is related to the variety and
can vary from white, to yellow, to orange, to red, to purple, to
brown, to black, to variations of the color range within the cob.
Wheat (Triticum aestivum) (Table 1(c))
Wheat is grown in all the temperate climates of the world and
is second to rice in terms of human consumption. True winter
wheat varieties require a period of cold to initiate the repro-
ductive stage of development (vernalization). Spring wheat
varieties do not need a cold period for ear formation although
they can respond to cold by flowering earlier. Wheat has a
higher protein content than any other major cereal, and a
further distinction is made in the type of flour produced.
Hard milling varieties produce flour suitable for bread produc-
tion. Soft milling varieties are used in the production of com-
modities such as animal feed or biscuits. Because winter wheat
has a longer growing season, they are usually denser in ear
formation than spring wheat (Photo 8). The average height of
wheat is 75150 cm, and unlike other cereal species, the stems
can be solid and hollow (Photo 9).
The wheat ear is made up of a series of spikelets arranged
alternately on the central rachis and each one is made up of a
cluster of six florets (Figure 8). Close study will identify the
different structures that form the spikelet, the most important
being the lower glume, which lies opposite the lemma with the
 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 59

Tassle Silk

Photo 6 Tassel and silk of maize.

Lax Medium Dense

(profile view)

Photo 8 Ear density wheat.

longer beak. In order to ensure consistency when assessing

Photo 7 Maize: structure of a corn cob.
their morphological characters, only the lower glumes from
spikelets in the mid third of the ear are examined in detail
(Figure 9). On studying the exterior surface of the glume
(dorsal view), distinct differences can be observed between
varieties. These include the length and curvature of the lower
glume beak and the width and shape of the lower glume
shoulder.
60 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties

The wheat grain can be separated by shape into those that
are round, elliptical, or ovate. At the opposite end to the
embryo, brush hair may be identified. If present, a distinction
can be made as to whether the hair is short or long in length
(Figure 10).
Durum Wheat T. turgidum (syn T. durum) (Table 1(d))
Commonly known as durum wheat and developed through
artificial selection many thousands of years ago, it is the
hardest milling variety of all the wheat species and used
extensively in the production of pasta. It grows well in coun-
tries bordering the Mediterranean where precipitation is low
and temperatures high. Similar in height (1.21.5 m) and
appearance to bearded wheat, but usually with a shorter
denser ear (Photo 10), durum wheat is characterized by
having well-developed, firm glumes and a pronounced keel
along the glume length (Figure 11(a)). The lemmas have a
rounded appearance due to the absence of a keel. The yel-
lowish, brown-colored caryopsis is often ridged. Brush hairs
are either absent or very short (Figure 11(b)).

Triticale (Table 1(e))

Triticale X secale was developed as a species in the nineteenth
century by crossing rye with wheat. It can grow to an average
height of 75150 cm, but does not tiller as much as wheat
and is used mainly as a fodder product. However, it has the
potential to produce a yield similar to that of some wheat and
Sections through stems of wheat straw. (cut midway between
ear and first node)
can be grown in poorer conditions more suitable for rye
production. Similar in height and density to wheat, the apical
Photo 9 Straw wall thickness. spikelet is sterile (Photo 11). At the top of the stem just below

(e)
Floret 6
Floret 5
Floret 4

(d ii)
(d i)
Floret 1
(c ii) Floret 2
(c i)

(a i) lower glume: always opposite (b i)

the lemma with the longer lemma beak (b ii)
(a ii) upper glume
(b i) lower lemma
(a ii)
(b ii) upper lemma
(a i)
(c i) palea
(c ii) palea
(d i) lemma - wheat caryopsis
between lemma and palea
(d ii) lemma
(e) apical spike
(f) rachis

(f)

Figure 8 Structure of a wheat spikelet.

 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 61

(a)
Extent of glume beak

Extent of shoulder
(b)

(d)

(e)

(f)

(c)

(a) lower glume beak

(b) shoulder
(c) keel
(d) papery membrane not used in
shoulder width assessment
(e) narrow wing
(f) broad wing (g)
(g) basal hairs
Figure 9 Structure of the lower glume.

brush hair long

brush hair short

Round Ovate Elliptical

Figure 10 Wheat grain, shape, and brush hair.
62 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
Photo 10 Triticum turgidum (syn T. durum).
the first rachis, it is possible in some varieties to observe neck
hairs. Glumes are of a thin papery structure and long and
narrow in shape. They may have hairs or weak toothlike
structures along the lemma margins (Figure 12(a)). In com-
mon with durum and rye, the glumes are strongly keeled
and have a much reduced shoulder. The elliptical caryopsis
is gray-brown in color and often ridged with a prominent
embryo. Brush hairs are present and short in length
(Figure 12(b)).
Rye (Secale X Secale) (Table 1(f))
Rye is a species that has been in cultivation for many centuries
and is the main cereal for bread production in parts of North
and Eastern Europe as it can withstand the cold better than
wheat and tolerate poorer growing conditions. It is also an
important forage crop. Depending on whether it is grown as
a fodder crop or for human consumption, rye will reach an
average height of 1.21.8 m. Until the caryopsis develops, the
ear has a narrow flattened appearance, and then, as the ear fills,
the caryopsis becomes highly visible between the lemma and
palea (Photo 12). The lemmas are strongly keeled and have a
series of toothlike structures along the margins and keel
(Figure 13(a)). The papery glumes are very narrow and have
no broad wing. The elliptical caryopsis is greenish brown and
strongly ridged and has a prominent embryo. Brush hairs are
present and of medium length (Figure 13(b)).
Barley (Hordeum vulgare) (Table 1(g))
Barley was one of the first cereal species to be cultivated, and
the grain has a wide variety of uses, from providing staple
food for humans and livestock to the production of certain
forms of alcohol. It is grown widely across the more tem-
perate zones of the world and ranks fourth in terms of food
production.
As a species, barley has more morphological characters that
are useful in determining varietal identity than any other
cereal. Depending on their tolerance to winter conditions and
whether they need a period of vernalization, barley varieties are
also classified as either winter or spring types. Winter barley in
particular has small straight hairs at right angles to the stem
(basal leaf sheath hairs) on the lowermost leaves (Figure 14). It
is important to observe the character at the optimum growth
stage (five to nine tillers) as assessment is more difficult once
the lower leaves wither and die.
In common with other cereals, the presence of pigment is a
major character in barley variety identification. Levels of pig-
ment vary in intensity and can be subject to environmental
factors such as sunlight and drought (Photo 13).
An ear of barley is classified as being a 6-row/4-row or
2-row, depending on whether each potential grain site has
developed and whether the developed grain has the character-
istic awn present (Photo 14). The size and shape of a barley
grain are determined by variety and environmental influences,
although six-row barleys have small asymmetrical lateral grains
caused by their proximity to the median grain. A conventional
two-row barley has two rows of undeveloped grain running up
through the center of the ear (sterile spikelets), either parallel
to each other or diverging out from the center. Where sterile
spikelets are so reduced in size to be barely visible, they are
classified as being deficiens (Photo 15). The grains sit on a
rachis with the first rachis segment located in a collar-like
structure at the top of the stem neck (Photo 16); the width,
length, and curvature of the collar provide further identifying
characters. The uppermost part of the stem immediately below
the collar is the neck.
Once the grain has been fertilized, the ventral surface of the
developing grain is protected by the palea, while the lemma
protects the dorsal surface and overlaps the palea on either
side. Sometimes, the lemma and palea do not meet properly
at the sides (gape), and it is possible to see the kernel within
(Figure 15).
Barley grains have two distinct views: the ventral view that
can be distinguished by the ventral furrow running the length
of the grain and the dorsal view that presents a number of
ridges (lemma nerves) on its surface (Figures 15 and 16,
respectively). Located on the dorsal side are the two-side
(lateral) lemma nerves, which extend down and form the
margins of the awn and the median (central) nerve. If pigment
is present, it will be evident in these nerves. On some varieties,
small thornlike structures (spicules) will be present. Spicules
occur frequently on the margins of the awns and sometimes on
the inner lateral nerves. They can be sparse, one or two per
nerve, or numerous and can extend out from the lateral nerves
onto the surface of the lemma on the lateral nerves at the awn
end (Photo 17).
 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 63

(a) glumes (narrow shoulder)

(b) lemma (not keeled)
apical floret present
Note: short ear, long awns

(b)

(b)

(a)
(a)

(a)

Caryopsis
Glume
elliptical, often
long and narrow ridged, brush hairs
firm, strongly keeled short or absent,
throughout yellow/brown in
color

(b)

Figure 11 (a) Triticum turgidum (syn T. durum) spikelet. (b) Triticum turgidum, glume and caryopsis.
64 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties

Note hairs on stem neck

Photo 11 Triticale X secale. Photo 12 Secale X secale (rye).

(a) glumes; long and narrow, thin

and papery, strongly keeled
throughout, glume shoulder much
reduced
(b) lemma; slightly keeled, may have
hairs and weak teeth along lemma
margin and keel

(a)
(b)
Glume Caryopsis
Note: strongy keel, may have Elliptical, often ridged, brush
weak hairs hairs present. (a) prominent
(b) embryo greyish brown in color
(a)

(a)

Figure 12 (a) Spikelet of Triticale X Triticosecale. (b) Triticale X Triticosecale.

 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 65

(a) glume; very narrow and

papery, no broad wing
(b) lemma, strongy keeled, strong
teeth along margins and keel

(b)

(a)

(a)

Caryopsis

Glumes elliptical, strongly

ridged, brush hairs
very narrow papery, no
present, greenish brown
broad wing
color
(a) prominent embryo

(b) (a)

Figure 13 (a) Spikelet of rye (Secale X secale). (b) Rye (Secale X secale).
66 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties

(b)

(a)
(a) basal leaf sheath hair
(b) hairs on base of leaf
blade

Figure 14 Barley plant showing basal leaf sheath hairs, and base of leaf blade.
 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 67

At the base of the grain and lying within the ventral furrow is observed on the dorsal side at the awn end, where it is
is a small rodlike structure covered in hairs (rachilla) and used possible to identify very small protrusions that create a rough
extensively in the identity of varieties. In some varieties, there surface on the areas between the lateral and median nerves
are an additional three distinct linked characters to observe in (sharkskin). It should be noted that this area may also be
barley grain. It is rare for these characters to be separated, so if associated with many rows of spicules.
one is present, then the other two will also be present. They are
fence hairs (ventral view) that lie in a vertical position at the
base of the grain and ventral furrow hairs that are extremely
small white hairs lying along the edge of the ventral furrow and
often extending beyond the rachilla. The third linked character

2 row 6 row

Photo 14 Ears of barley showing different structures between a two-

Photo 13 Range of pigmentation in different varieties of barley. row barley and a six-row barley.

Deficiens Divergent Parallel

Photo 15 Barley sterile spikelet attitude in two-row varieties.
68 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties

Platform collar Cup collar

Photo 16 Barley first rachis segment and collar.

(e)

(a) lemma
(b) palea
(c) ventral furrow
(d) lemma base (attachment
point to rachis)
(e) broken off awn
(f) rachilla
(c)
(g) gape may not always be
present
(h) inter glume hair

(a)

(b)

(g)

(f)

(h)
(d)
Figure 15 Ventral view of grain of barley.
 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 69

(a)

(b)

(g)

(c)

(d)
(d)

(a) original attachment to rachis

(b) embryo
(c) median nerve
(d) lateral nerves
(e) outer lateral nerves (note (h)
spicules)
(f) broken off awn
(g) lemma (e)
(h) area where sharskin is
observed (if present) (f)

Figure 16 Structure of a barley grain dorsal view.

70 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties
Photo 17 Barley: dorsal view showing spicules on inner lateral nerves.
Equilateral Unilateral
(spreading) (one sided)
Photo 18 Oat panicles.
Oats (Avena sativa) (Table 1(h))
Oats are an important crop providing food for human con-
sumption and livestock and are particularly useful in NW
Europe where they tolerate the cooler and often moister con-
ditions during the growing season. On average, an oat plant
reaches a height of 1.21.8 m, but oat characters such as height
and the frequency of awns are liable to be affected by environ-
mental factors. Hairs on the basal leaf sheaths are present in
some varieties (Figure 14) and on the margins of the leaf
blade. They are best seen at leaf emergence when they vary
from a few at the base to half the leaf length.
The uppermost node (upper culm node) from which the
flag leaf emerges is a swollen thickening of the stem and
shrinks on maturity. Fine hairs above and below the node
are present in some varieties. The branches of the panicle
emerge from swollen nodes on the flowering stem with single
spikelets interspersed along the main branches (Photo 18).
A single spikelet will develop into one (primary) that is
always the largest grain and can develop into second (sec-
ondary) and third (tertiary) grains, each attached to the other
by a rachilla.
Each caryopsis is protected by a lemma and a palea, the
whole protected by papery glumes (Figure 17). Awns if present
emerge from the median nerve of the lemma of the primary
grain. At the base of each grain, grain basal callus hair may be
present at either side. The rachilla can vary in length from short
to long and vary from plain to strongly grooved.
Conclusion
From the earliest times, man recognized the different cereals as
an important food source. The crops they produced were sub-
ject to continual change and selection over the decades result-
ing in the gradual improvement of yield and quality. At the
same time, others were recognizing similarities between plants
and evolving a system that placed them into taxonomic groups.
The more detailed and precise the study of plants and their
morphology became, the more demand it placed on using a
greater range of identifying morphological characters. It was
this assessment of morphological characters that gave the dis-
criminatory evidence essential for laying the foundation for
varietal identification, which could in turn be used for estab-
lishing variety ownership through the introduction of plant
variety rights. Modern techniques such as the use of DNA
fingerprinting are already being used to determine varietal
identity, but for the purposes of UPOV and plant breeders
rights, they can only be used as an addition to the use of
morphological characters.
Exercises for Revision
Using the UPOV technical guidelines for a cereal species of
major importance within your country, identify the charac-
ters that could be used to generate a description for the
most widely grown variety.
Select a species of lesser importance and write a description
for a variety based on the UPOV guidelines for the species.
Using the UPOV guidelines, select one of the species grown
widely within your country and study in detail those char-
acteristics not covered in this chapter. Write a summary as
to how these additional characters could be used to pro-
duce a detailed description of a variety, illustrating with
photos or drawings where necessary.
A characteristic of cereal species is the presence of glumes.
Write a paper on the importance and evolution of the
glume to each.
 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 71

Spikelet structure

(a) glumes
(b) primary grain
(a)
(c) secondary grain
(d) awn
(b)
(e) basal callus hair present, 1-2 per
side of in tufts
(f) lemma
(g) palea
(h i) rachilla (note length)
(h ii) rachilla medium long in length
(i) basal callus (d) (c)

(i)
(e) (i)

Primary grain characters

(h i)

(h
ii)
(g) (f) (f) (g)

Figure 17 Oat spikelet and grains showing spikelet structure.

Exercises for Readers to Explore the Topic Further
Currently, variety descriptions are produced using methods
that have remained largely unchanged since their introduc-
tion. Some biotechnological methods have now been
incorporated into existing systems. Using a species with
which you are familiar, identify any available biotechnol-
ogy methods and undertake research to assess the advan-
tages that could be gained by their inclusion.
Could varietal identification be carried out partly or wholly
by biotechnology?
See also: Grain Marketing and Grading: Identification of Varieties
of Food Grains; The Basics: Grain: Morphology of Internal Structure;
Taxonomic Classification of Grain Species; Wheat-Based Foods:
Cereals: Overview of Uses: Accent on Wheat Grain.
Further Reading
Detailed descriptions of characters for varietal identification can be found in the UPOV
Guidelines under the following headings: (upov.int/test guidelines/choice of
language). The guidelines are available worldwide and provide a standardized
format for the characters used in varietal identification.
72 THE BASICS | Grain and Plant Morphology of Cereals and how characters can be used to identify varieties

Barley (Hordeum vulgare) UPOV TG/19/10. NIAB: Botanical descriptions of varieties info@niab.com.
Durum wheat (Tritium turgidum syn T. durum) UPOV TG/120/3. Oat (Avena sativa) UPOV TG 20/10.
Hervey-Murray CG (1980) The Identification of Cereal Varieties. Cambridge: Cambridge Rice (Oryza sativa) UPOV TG16/8.
University Press. Rye (Secale x secale) UPOV TG58/6.
HGCA: Milling wheat: quality control criteria and tests. The biology of maize www.ogtr.gov.au.
Ikisan: morphology of rice. The biology of Triticum turgidum www.inspection.gc.ca/triticum turgidum.
Ikisan: morphology of maize. Triticale (triticale) X triticosecale UPOV TG/3/11.
Institute of rice research www.irri.org. Wheat (Triticum aestivum) UPOV TG/3/11.
Maize (Zea mays) UPOV TG2/7.
 The Cereal Grains

Contents
An Overview of the Family of Cereal Grains Prominent in World Agriculture
Cereals: Domestication of the Cereal Grains
Maize: Overview
Wheat: An Overview of the Grain That Provides Our Daily Bread
Durum Wheat: Overview
Rice: Overview
Wildrice, Zizania: Overview
African Rice (Oryza glaberrima): A Brief History and Its Growing Importance in Current Rice Breeding Efforts
Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses
Sorghum Grain, Its Production and Uses: Overview
Rye Grain: Its Genetics, Production, and Utilization
Triticale: Overview
Oats: Overview
Coix: Overview
Millet Pearl: Overview
Millet Minor: Overview
Teff: Overview

An Overview of the Family of Cereal Grains Prominent in World Agriculture

CW Wrigley, The University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.

Topic Highlights Introduction

Cereal grains are produced by flowering plants of the grass
family.
Wheat, rice, and maize (corn) are the major cereals, each
produced in quantities of about 600 million tonnes annually.
Other cereals are barley, rye, triticale, oats, sorghum, and
the millets.
Cereal grains have a high proportion of starch, present as
distinct granules in the floury endosperm.
Cereals are a valuable source of protein; their low fat con-
tent is mainly in the germ (embryo), from which the new
plant emerges during germination.
Processed foods from cereals include a wide range of bread
types, pasta, noodles, breakfast cereals, beer, and porridge.
Cereals are major sources of animal feed (grain and plant)
and fuel for transport.
Learning Objective
To achieve understanding of the agriculturally significant
species and uses of cereal-grain family.
Colloquially, we often use the word cereal to mean a breakfast
cereal, such as cornflakes. The manufacture of cereal in this
narrow sense is described in this encyclopedia, but more
broadly, the term cereal applies to the many grain genera
that are flowering plants of the grass family (Poaceae or Gra-
mineae), whose seeds are used as food. It may be difficult to
imagine wheat, for example, as a flowering plant, but wheat
flowers are illustrated in Figure 1. (And do not get confused
between this flower and the flour of similar-sounding name,
meaning crushed grain.)
Seed or Grain?
The name cereal derives from Ceres, the Roman goddess of
grain. The words seed, kernel, and caryopsis are also used to
denote cereal grain. In addition, the term corn is sometimes
used for the grains of cereal species in general, but more often
(especially in the United States), the word corn denotes the
cereal species maize, also known as Indian corn. Wheat and
barley seeds are shown in Figures 2 and 3, as they appear by
scanning electron microscopy.

Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00006-1 73

74 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture

Figure 1 A flower of wheat. The outer glumes have opened wide.

Anthers are seen above the feathery style, which will receive pollen grains
shed by the anthers.

Cereal grain serves two contrasting functions:

1. For the plant, the mature grain is solely of significance as a

seed, that is, the means by which the species is perpetu-
ated. When the plant has died, the ongoing life of a cereal
seed is in its embryo or germ. Most of the grains mass is
taken up by the endosperm (Figure 4), which is the storage
organ of the grain. It contains the starch and protein for the
rainy day when moisture will trigger the dormant seed to
start swelling and producing roots and a shoot. In that
situation, the endosperm provides the essential nutrients
for germination before the emergence of the green leaf,
allowing photosynthesis to take over as the source of energy
and nutrients.
2. For mankind, the mature grain is mainly of significance
as an important source of nutrients, that is, an item of
food for humans and a source of feed for mans animals.
Examples of these many processed foods are shown in
Figure 5.
The major cereal grains of economic importance (Figure 6) are
the cool-season crops (wheat, barley, oats, and rye) and the
warm-season cereals (rice (paddy), maize (corn), sorghum,
and the millets). Table 1 lists their botanical names and
major uses. Cereals are monocotyledonous plants, as distinct
from the dicotyledonous members of the wider family of
Figure 2 The wheat grain (a) in dorsal view and (b) in ventral view,
showing the brush (top left) at the end opposite from the point of
attachment (the hilum) of the grain to the rachis of the wheat head.
The crease extends along the ventral side of the grain. The germ can
be seen as a rough area in the dorsal view (a) at the hilum end. These
images were provided by scanning electron microscopy, with the
grain resting on the stage of the microscope. Reproduced from Wrigley
CW (2004) Cereal sprouting. In: Goodman RM (ed.) Encyclopedia of
Plant and Crop Science. New York: Marcel Dekker, with permission.
grains. The dicot grain species include the oilseeds and the
pulses (grain legumes).
Cereal plants are also divided according to their photosyn-
thetic systems, namely, either C3 or C4. The C4 photosynthetic
carbon cycle offers metabolic advantages over C3 photosyn-
thesis in dry situations with high light intensity and high
temperature. The approximately half of grass species that are
C4 include maize, sorghum, and millet.
 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture 75

Figure 3 Scanning electron micrographs of (a) a barley grain, covered by the integrated glumes (lemma and palea); (b) the rachilla lying in the
groove of the crease; (c) the base of the rachilla with many of the long hairs broken off as a result of grain handling; (d) dust from the movement
of barley grain showing the spear-like rachilla hairs. Note the different degrees of magnifications: the black bar at the bottom right represents a length of
(a) 1 mm, (b) 400 mm, (c) 40 mm, and (d) 100 mm.

Historical Perspective
Cereal grains have the great distinction of being the catalyst
that transformed humans, as the hunter-gatherers into agricul-
turalists no longer living as nomads. The origins of civilization
involved the enormous step of primitive mankind discovering
that it was possible to remain in one place where cereal seed,
sown intentionally, would yield a reasonably reliable source of
food, which could be stored indefinitely. As a result, people
found that a fixed dwelling place could be established near
cereal crops. Further consequences were the development of
permanent forms of housing, the domestication of animals
(fed with cereal grain), and the spare time to develop the
many other characteristics of civilization.
Early consumption of cereals involved merely chewing the
raw grain. Primitive forms of processing were developed, such as
grinding the grain between stones and soaking the grain in water
to make it easier to chew. Next, heat was discovered as a further
means of easing mastication and digestion, producing a porridge
or gruel. Placing a stiff porridge in the hot stones or ashes of a fire
would have made it more palatable, resulting in baked foods.
A porridge left overnight would gain wild yeasts, leading to
the discovery of leavening as a means of making the baked
product lighter. Further refinements would have been the siev-
ing step, to remove husks and bran particles after grinding, and
the identification of wheat as the ideal grain for the baking of
leavened bread due to its dough-forming properties.
There is archaeological evidence that primitive baked prod-
ucts were being made in the late stages of the Stone Age.
Purpose-built ovens were used in ancient Egypt as far back as
2700 BC, with reasonably sophisticated milling and baking a
millennium later. As a result of this long history of cereal-grain
use, diverse uses have been developed worldwide for their
processing, taking advantage of the specific characteristics of
the individual cereal species (see Table 1 and Figures 5 and 7).
World Production of Cereals
Cereal grains are grown extensively in all continents, with the
obvious exception of Antarctica. They are the major sources of
food for mankind, especially in developing countries. In
76 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture

Brush

Aleurone
cells

Cuticle Storage
protein
Epidermis
Hypodermis Cell nucleus
Cell wall
Thin-walled
cells
Endosperm
Cross cells
Testa Starch
granules
Nucellar
epidermis Figure 5 Some of the many foods made from cereals (mainly wheat).

Pericarp Aleurone grains

Scutellum
Plumule
Embryo
Root
(a) Sheath and cap
(b)
Figure 4 (a) Diagram of the many parts of a wheat grain, cut in half
to show its subcellular structure. (b) Scanning electron microscope
image of the cut surface of a wheat grain. The bran layer, visible at the top
left, covers the layer of square-shaped aleurone cells, which in turn
enclose the endosperm (the lower right half of the photo). Starch
granules can be seen as flattened spheres embedded in the protein
matrix, surrounded by endosperm cell walls.
addition to their use as a major source of feed for animals,
cereal grains are extensively processed industrially for food and
nonfood products.
Table 2 provides grain-production statistics for the worlds
cereal-growing countries. The statisticians term coarse grains
includes maize, barley, rye, triticale, and oats. In this case, the
production statistics for rice appear much less than in some
comparable tables because they show volumes for milled rice,
with the outer coverings removed.
The provision of such data for one harvest year permits
direct comparison between cereals and countries, but it
should be regarded as a snapshot of production and trade,
because of the fluctuations that occur from 1 year to another.
Figure 8, on the other hand, shows how statistics vary from
year to year for all cereals, for their production and utiliza-
tion, with the added indications of carryover stocks from year
to year. The statistics in Table 2 also illustrate the regional
distribution of centers of production for the prominent
cereals, namely, tropical countries for rice (and maize) and
cooler regions for wheat and other coarse grains.
The annual world production of all cereal grains is now about
2.5 billion tonnes (Table 2, Figure 8). Assuming a world
population of about seven billion people, this amount of grain
represents about 360 kg of cereal grain per person per year or
1 kg per person per day for the whole worlds population. It thus
appears that world hunger could readily be alleviated, assuming
that all cereal grains might be used for food and assuming that it
could be transported to all the regions where it is needed. Of
course, this is neither practicable nor possible, since many of the
regions of production are distant from the places of need. In
addition, the worlds animal agriculture and industrial opera-
tions are reliant on significant amounts of cereal grain.
World production of wheat has ranged between 690 and
710 million tonnes (Mt) annually in recent years. This produc-
tion has come from an area of about 220 Mha, with a world
average yield of 3.2 tonnes ha 1. Two-thirds of world produc-
tion of wheat goes to food; the remainder goes to a combina-
tion of seed use, animal feed, and industrial processing. World
trade in wheat is about 20% of total production.
Production of wheats close relative, rye, is minor by com-
parison with wheat; world rye production is about 20 million
tonnes annually, the main production regions being in eastern
parts of Europe, especially Poland, Germany, and Russia.
World production of rice is similar to that of wheat, if the
production of unmilled paddy rice is considered. On the basis
 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture 77

Figure 6 Grains of major cereal species: (a) bread wheat; (b) durum wheat; (c) triticale; (d) rye; (e) barley; (f) oats; (g) sorghum; (h) corn; (i) Japanese
millet. Adapted from Fitzsimmons RW and Wrigley CW (1984) Australian Barleys: Identification of Varieties, Grain Defects and Foreign Seeds, 2nd edn.,
pp. A11A13. Melbourne: CSIRO.

of milled rice, annual world production averages over 500
million tonnes. International trade in rice is much less than
for wheat and maize. Rice, together with maize, provides the
major energy and protein source for many cultures.
Annual production of maize (950 million tonnes in 2013/
14) is greater than that of wheat. Statistically, maize is included
in coarse grains, production of which totals 1.2 billion tonnes
annually (Table 2). World trade in coarse grains is about 100
million tonnes. In addition to maize, coarse grains include barley
(production averaging about 150 million tonnes annually), sor-
ghum (50 million tonnes), and oats (30 million tonnes).
Morphology of Cereals: What They Look Like
Cereal grasses have narrow leaves, hollow jointed stems, and
spikes or clusters of membranous flowers (Figure 1). Figure 9
shows how the mature grain of wheat is held within glumes on
78 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture

Table 1 Cereal-grain species (in taxonomic groups) and their uses

Common
names Genus and species Uses

Pooids
Common Triticum aestivum Breads (leavened and unleavened), noodles, breakfast cereals (processed porridge), cakes, cookies,
wheat chapatis, snack foods, bulgur, ingredients for a wide range of foods, animal feed, industrial uses
Durum Triticum turgidum Pasta (e.g., spaghetti and macaroni)
wheat subsp. durum
Rye Secale cereale Bread (often together with wheat), crispbread, spirits
Triticale Triticosecale Bread (often together with wheat), animal feed
Barley Hordeum vulgare Malt, beer, spirits, porridge, cooked grains (e.g., soup ingredient)
Oats Avena sativa Porridge (as rolled oats, oatmeal), muesli, oatcakes, animal feed
Oryzoids
Rice Oryza sativa Cooked and fried grain, sake, extruded snack foods
Wild rice Zizania aquatica Cooked grain
Chloridoids
Teff, Indian Eragrostis tef Cooked grain, porridge
millet
Finger millet Eleusine coracana Cooked grain, porridge
Panicoids
Corn Zea mays Polenta, breakfast cereals, tortilla, cornbread, porridge, snack foods, a wide range of industrial uses
(maize) (e.g., corn syrup and food ingredients)
Sorghum Sorghum bicolor Animal feed, cooked grain, porridge, beer, flat cakes
Pearl millet Pennisetum glaucum Cooked grain, porridge
Common Panicum miliaceum Cooked grain, porridge
millet
Japanese Echinochloa species Cooked grain, porridge
millet
Foxtail millet Setaria italica Cooked grain, porridge

The flowers of oats and rice, on the other hand, are arranged
in an inflorescence called a panicle (Figure 11). The grains of
oats and rice do not thresh as the free seed. The term groat
applies to the oat seed with the glumes removed. The term
paddy rice, or just paddy, refers to the rice grain with its
husks in place, the form in which it is harvested. The familiar
cob (ear) of corn (maize) is distinct from the seed-bearing
structures of the other cereals described earlier (Figure 12).

The Diversity of Cereal Grains

Figure 7 Chinese steamed bread.
the single stem of the head (ear) (Figure 10). The threshing
process at harvest separates the wheat grain (as in Figures 2
and 6) from the glumes, rachis, and other parts of the plant.
The structure of the barley head (Figure 10) is similar to that of
wheat, but on threshing, the glumes of barley remain with the
grain (Figures 3 and 6), whereas the wheat grain threshes free
from the glumes. The rye grain also threshes free of the glumes
(Figure 6).
Wheat
Wheat is prominent among cereals, because of its unique
dough-forming properties. These viscoelastic characteristics
are due to the glutenprotein complex, formed from the
major storage protein of its endosperm. This is the basis of
mans attraction to wheat the reason for the annual cultiva-
tion of over 1014 wheat plants because wheat gluten alone
can sustain mans desire for leavened bread products.
There is a great diversity of uses of common or bread wheat
(Table 1), such as pan breads, pocket (Arabic) breads, noodles,
and steamed breads (Figures 5 and 7). In addition, there is a
diversity of processing methods, especially for the range of
leavened breads. There is thus the need to breed a range of
wheat varieties to suit the various processing specifications. In
addition, there is the need for the appropriate variety to be
grown in a suitable climate and region, with the right manage-
ment, aimed at achieving target attributes of bulk density, lack
 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture 79

Table 2 World production in the year 2013/14 of cereal grains by countries/regions, in millions of tonnes (metric tons)

Region Wheat Coarse grainsa Riceb All cereal grains

Argentina 11 35 1 47
Australia 26 11 1 37
Brazil 5 73 8 86
Canada 33 26 59
China 121 218 142 480
EU 143 159 2 304
India 93 42 105 240
Indonesia 9 38 47
Mexico 3 29 33
Pakistan 24 4 6 34
Russia 52 36 1 88
South Africa 2 14 15
Thailand 5 21 26
Turkey 18 13 1 32
Ukraine 22 38 60
The United States 58 372 6 436
World 706 1252 473 2432
a
The category coarse grains includes maize, barley, rye, triticale, and oats.
b
Calculated as milled rice (not paddy).
Source: Data from United States Department of Agriculture.

Cereal production, utilization and stocks

Million tonnes Million tonnes

2500 800

2300 600

2100 400
Awn

1900 200 Palea

Grain
1700 0
2003/04 2005/06 2007/08 2009/10 2011/12 2013/14
fcast Lemma

Production (left axis) Utilization (left axis) Outer glume

Stocks (right axis) Rachis

Figure 8 Production, utilization, and stocks of cereal grains during

recent years. Reproduced with permission from FAO.
of defects and contaminants, grain hardness, protein content,
milling quality, and dough-forming properties. It is usual to
segregate wheat after harvest into specific grades, each being
suited to appropriate uses and prices. These grades carry
through into world trade, forming a basis for buyers to select
based on the combination of quality and price.
Durum, a distinct species of wheat (Table 1, Figure 6), has a
harder, larger grain, with a yellowish hue (amber according to
the title of the international durum grade). Because its endo-
sperm is so hard, it fragments on milling into particles that are
larger than the particles from common wheat (especially soft
Figure 9 A spikelet of wheat. The mature grain is held within glumes,
and all are attached to the rachis (the central stem of the head). Other
spikelets have been removed from the rachis for clarity. Reproduced
from Fitzsimmons RW, Martin RH, and Wrigley CW (1983) Australian
Wheat Varieties: Identification According to Growth, Head and Grain
Characteristics, 2nd edn., p. 13. Melbourne: CSIRO, with permission.
wheats). This coarse flour from durum wheat (semolina) is
suited for pasta manufacture (extruded from a relatively dry
dough) and also for couscous, frike, and bulgar (also known as
bulghur or burghul). Some of these foods are also made
from common wheat in some regions.
80 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture

(a) (b) (c)

Figure 10 (a) Heads of wheat. (b) Heads of two-row barley. (c) Heads of six-row barley.

Spikelets

Branch

Rachis

Figure 12 The cob of corn.

First node of rachis

with whorl of branches Rice

Neck Like wheat, rice is another cereal with a special place in mans
diet, providing a dietary staple for over half of the worlds pop-
Flagleaf ulation, thus constituting their major source of energy and pro-
Top ligule tein. Unlike other cereals, rice is usually consumed as the whole
grain, after removal of the bran layers. Rice can be grown under a
wide range of climates, either in flooded irrigation or under
Flagleaf
sheath dryland conditions. Some 90% of the worlds rice production is
Uppermost
stem node in Asia, especially in developing countries. International trade in
rice accounts for only about 4% of world production.
Prices vary greatly depending on the type of rice. For exam-
ple, Basmati scented rice from Pakistan and northwest India
Figure 11 The head (panicle) of oats. Reproduced from Fitzsimmons RW, might command four times the domestic price of ordinary
Roberts GL, and Wrigley CW (1983) Australian Oat Varieties: Identification of rice. Quality grades of rice are partly based on dimensions
Plants, Panicles and Grains, p. 11. Melbourne: CSIRO, with permission. long-, medium-, and short-grain rice. Quality attributes relate
 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture
Figure 13 Rice-based fast food sold by the road side in Thailand,
demonstrated by Professor Siriamornpun of Mahasarakham University,
Thailand. Rice has been cooked inside a length of bamboo. Stripping off
the outside of the bamboo reveals the cooked rice inside.
to the milled grain: shape, color, translucency, and uniformity
and also the absence of broken grains.
For human consumption, paddy rice is dehulled and milled
to remove the bran layers, leaving the familiar lustrous, white
grain. Brown rice, much less popular for eating, retains the
bran layers, together with added nutrients. Short-grain waxy
(having low amylose content) varieties (japonica types) usually
become sticky on cooking, a characteristic that is preferred in
many parts of north Asia. There is a preference in many
Western countries for the drier, flaky quality usually provided
by the long-grain nonwaxy varieties (indica types), more often
grown in tropical regions.
A novel example of rice for food is illustrated by the Thai
fast food in Figure 13. Rice, water, and flavoring have been
cooked inside the stem of a length of bamboo. The result is
sold by the roadside. To eat the cooked rice, the outside of the
bamboo is stripped back; the cooked rice is retained by the
membrane inside the bamboo.
Whereas milled rice is mainly consumed as the cooked grain,
there are also many food products made from rice, namely,
parboiled rice, rice crackers and noodles, rice cakes and snack
foods, rice flour, and fermented drinks (sake and rice wine).
Maize (Corn)
Maize, indigenous to the Americas, is produced worldwide
mainly in tropical and warm regions. It is an annual plant with
81
Figure 14 Plants of maize (corn). The pollen-dispersing tassels are
at the top of the plants. Developing cobs, sheathed in leaf material,
appear halfway up the stalk of the plant on the right.
an erect, leafy stalk. The ear consists of a central pithy section
(the cob, Figure 12), via which nutrients are transported from
the leaves and stalk to the grains (some hundreds or up to one
thousand grains per cob). The tassels (Figure 14) of the imma-
ture cob constitute the inflorescence bearing the male flowers.
There is a wide range of types of grain. Variations in grain
types include size, color, and hardness. Colors range from
white, to yellow, to red, to almost black. Endosperm types
include soft and floury to hard and flinty. Popcorn is a special
type of endosperm that expands suddenly on heating, due to
vapor expansion inside the grain. Sweet corn, with an unusually
high sugar content, is generally harvested immature, retaining a
high water content to provide the familiar juicy mouthfeel.
There is extensive industrial processing of maize, especially
in the United States and Europe. An important group of prod-
ucts is the wide range of breakfast cereals and snack foods.
Industrial processing also involves fractionation of the maize
grain by wet milling into its major biochemical components.
The main product of processing is cornstarch, which is the
largest component of the corn endosperm. By-products are
maize oil, from the germ and the protein fraction, which is
traded under the inappropriate term corn gluten. These direct
products may be further processed, especially the starch, much
of which is hydrolyzed to simple sugars as a sweetening agent,
as well as being used to make syrups and spirits.
Barley
The main uses of barley (Figures 3 and 6) are for animal feed
and for the production of pearled barley, malt, and beer
(Table 1). These two uses (feed and food) form the basis of
82 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture
the two main grades for trade in barley (known as feed barley
vs. malting barley). Another major distinction is based on head
morphology, namely, two row or six row depending on
whether there are grains in the full set of florets in the head
(six row) or if there is only one set of grains on either side of the
head, with the other florets sterile (Figure 10). In some barley-
growing countries (Europe and Australia), six-row barley vari-
eties are often of feed quality, but six-row varieties are also of
malting quality in North America.
Barley is the grain used for Scotch and Irish whiskies. In
addition, a proportion of the barley crop is used directly for
human consumption; after pearling (abrasion) to remove the
adhering lemma and palea, the barley grain is boiled in soup as
the whole pearled grain, or it is coarsely ground and cooked as
gruel. Barley flour is also used for a range of baked products,
but barley is not suitable for the production of leavened bread,
lacking the gluten proteins of dough-forming quality. Never-
theless, barley appears to have been used for bread making by
ancient civilizations. Barley, for example, appeared on Greek
coins several centuries BC, and it was the staple diet of Roman
gladiators, who were thus known as hordearii (reflecting the
origins of the genus name, Hordeum).
Oats
The oat grain and plant and grain are illustrated in Figures 6
and 11. Oat plants thrive in temperate climates, with cooler
moister conditions, compared to other cereals. Oats are often
used as fodder for stock, grazing off the growing plants. In such
cases, it may also be possible for the farmer to harvest a crop of
grain, which in turn may also be stored on-farm for stock feed.
In addition, oats may be grown primarily as a grain crop,
generally using varieties that have been bred specifically for
use as grain. A large proportion of harvested oats is used for
animal feed, especially for horses.
Compared to the other cereal grains, the oat grain tends
to have higher contents of protein, lipid, and soluble fiber
(b-glucan). Oats are reported to have beneficial hypocholester-
olemic properties and are useful in the management of the
insulin response in diabetics. For human consumption, the
adhering glumes are abraded from the oat grain. The resulting
groat is cut, rolled, or ground to yield products such as oatmeal,
rolled (flaked) oats, and oat flour, for processing into instant
oats for porridge, oat cakes, breakfast cereal, and infant foods.
The groat has a higher content of lipid (fats) than wheat, plus
significant lipase activity (fat-splitting enzyme), and these fac-
tors must be taken into account in oat processing. There is the
consequent risk that after milling, fat rancidity, bitterness, and a
soapy taste will be produced by the actions of various endoge-
nous enzymes, namely, lipase, lipoxygenase, and peroxidase. To
prevent this, it is usual for milled oats to undergo some form of
heat treatment to inactivate the lipase activity. This is often done
by steam treatment, together with kiln drying.
Rye and Triticale
Rye is the only cereal grain that approaches wheat with respect
to bread-making properties, although the achievement of
bread quality in rye breads usually requires the incorporation
of a considerable proportion of wheat flour (Table 1, Figure 6).
Rye is a popular human diet in Europe. The production of rye
in Germany exceeded that of wheat from about 1940 to 1960,
but more recently, the production of wheat has increased
considerably, and a higher proportion of the rye crop has
been used for animal feed.
Rye is particularly susceptible to contamination with ergot,
a toxic fungus that replaces the grain in the rye head during
development, so that a long black ergot body protrudes from
the mature head. Ideally, the production of ergot in the crop is
avoided, but there are techniques for the removal of intact
ergot bodies prior to milling.
In addition to the manufacture of rye breads, rye-based
crispbread is an important use of rye, generally with rye whole-
meal or from flaked rye. Other baked products include Ryvita
and pumpernickel. In human nutrition, the rye grain offers an
unusually high level of pentosans. This source of soluble fiber
offers advantages of giving a feeling of satisfaction, thus help-
ing in slimming diets, as well as reducing the rate of rise in the
blood sugar level after ingestion.
Triticale is a man-made crop that combines the genomes of
wheat and rye (Table 1). Most cultivated triticales are hexa-
ploid types, containing the genomes of durum wheat and rye.
It thus tends to be intermediate between rye and wheat in its
grain composition and characteristics. The expectations for the
grain have been that it would contribute the disease resistance
and growth hardiness of rye to wheat while also combining the
baking quality of wheat with the unique flavor of rye bread.
Triticale has thus been used for baking; nevertheless, its main
use is for animal feed.
Sorghum and the Millets
The grains of sorghum and of various millet species (Table 1,
Figure 6) constitute a major source of protein and energy for
many people in Africa and Asia, where these grains are used as
porridge, boiled in water after grinding. In such situations,
these cereals offer the advantage for subsistence farming of
tolerance to drought and suitability for tropical growth envi-
ronments. Pearl millet is the most widely grown of the millet
family, with an estimated area worldwide of about 30 Mha.
Second in importance are common millet and finger millet
(Table 1). Many millets, due to small grain size, are used as
feed for wild and domesticated birds.
In addition to the use of sorghum in cooking, it is also dry-
milled and fermented for various forms of beer. On the other
hand, the main use of sorghum in developed countries is for
animal feed and, to a lesser extent, for industrial processing.
There are various mutant forms of sorghum with distinct char-
acteristics that include high lysine content, waxy starch, high
sugar content (stalks), and even scented endosperm. The grain
of many sorghum varieties is pigmented, largely due to tannins
located in the outer layers (testa) of the grain. The high-tannin
varieties are also somewhat less nutritious, because the tannins
interfere with protein digestibility. Some sorghum varieties
have bitter flavors, which confer resistance to attack by birds.
Composition
Starch, a polymer of glucose, is the major component of the
endosperm of cereal grains (Table 3). It is the stored form of
 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture 83

Table 3 Approximate composition of the major cereal grains, as percentages of dry weight of whole grain, except for rice

Component Wheat Rice (milled) Maize Barley Oats (groats) Rye

Starch 7580 8590 7580 7580 7080 7580

Protein 916 811 1012 912 1215 1215
Fat 23 0.51.0 410 12 68 12
Crude fiber 23 0.20.6 23 56 12 23
Minerals 12 0.40.6 12 23 23 23

In addition, grains (as is) would contain moisture at a level of 915%.

energy that is released on germination when amylase enzymes

are produced to break the starch down to glucose units for the
developing embryo, roots, and shoots. For feed and food,
starch also provides the major source of energy, providing it
in a slow-release form that is well suited to our digestive
systems. Although cereal-grain proteins do not have the ideal
combination of essential amino acids, being slightly deficient
in lysine especially, they are generally ingested together with
other sources of protein whose amino acid composition com-
plements that of cereal grains. The relatively low fat content of
cereal grains is a dietary advantage, especially when coupled
with the levels of nonstarch polysaccharides, which act as
important sources of fiber in the human diet.

Cereal Grains and Our Health

For some individuals, cereal-based foods may cause dietary

problems. One of the best characterized of these intolerances
is celiac disease, a condition caused by the ingestion of wheat
glutenprotein and of related grain proteins, namely, rye, trit-
icale, barley, and (for some individuals) oats. Despite its mis-
leading name, buckwheat is not a grain that is toxic to celiacs.
On the other hand, cereal grains (milled or not) provide
good sources of protein, fiber, and complex carbohydrates,
earning for cereal grains a major place in dietary guidelines
(Figure 15). Additional health benefits may be provided by
the inclusion of the outer layers of cereal grains, in whole-grain-
foods. This is because these parts of the grain (Figure 4(a))
provide an additional source of vitamins and minerals and
because they contribute increased amounts of fiber to the diet.
During their growth, cereals also produce pollen that may
cause allergy problems on inhalation. The pollen grains of the
various cereal flowers have distinctive shapes (Figure 16). The
germination of the pollen grain commences on exposure to
moisture, normally when it alights on the stylus of a flower of
the same species (Figure 1), but the same reaction may occur if
the pollen comes in contact with the mucosa of the respiratory
tract. The proteins so released may cause swelling of the
mucosa, resulting in rhinitis and asthma. The cultivated cereals
may be the cause of this scenario, but it is more common for
inhalant allergy to be caused by a wider range of plants species,
not necessarily propagated for their grain.
Dust from cereal grains also causes irritation in the respira-
tory tract. Dust produced during the movement of barley grain
(Figure 3(d)) consists partly of hairs broken off the rachilla
(Figure 3(b) and 3(c)), which lies in the crease of the barley
grain. Barley varieties differ in the length of the rachilla hairs,
Figure 15 An example of the nutrition pyramid recommending that
foods containing grains (base of pyramid) should make a greater
contribution to our diet than other foods, such as fruits, dairy, meat,
and fats.
this characteristic being useful in visual variety identification.
Irritation of the skin and respiratory tract is worse for dust from
varieties with long rachilla hairs.
Future Prospects
Although world production of cereal grains is equivalent to
1 kg per person per day for the worlds population, many parts
of the world still experience hunger. How can we share the
cereal-grain harvest more equally? This problem is epitomized
in the inscription on the old plate in Figure 17; the translation
reads: Old bread is not hard no bread, that is hard.
With this old problem in mind, new research methods are
being directed toward improving the yield and quality of the
grain produced, especially by breeding. Despite the success of
cereal grains as the major food source for mankind, there are
many stresses that reduce grain yield. These stresses include a
range of pathogens and predators, as well as many abiotic
factors (heat, cold, frost, drought, waterlogging, and heavy-
metal toxicity).
Traditional breeding has provided great improvements and
it will continue to do so. These advances are being assisted by
the identification and characterization of specific remedial
genes. Some of these genetic opportunities lie in plants of
84 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture

Figure 16 Pollen grains from the flowers of cereal species, revealed by scanning electron microscopy.

species distantly related to the species of interest. Novel genetic
technologies (genetic modification (GM)) facilitate these cross
species transfers, thus augmenting conventional breeding,
leading to new varieties that are better at surviving stresses,
plus providing novel grain qualities.
GM technologies have been more successful in commercial
practice for noncereal species, especially soybean, cotton, and
canola. Nevertheless, GM corn is grown widely, especially in
the United States, accounting for about one-third of the area
worldwide for GM crops.
For other cereal species, GM and conventional breeding
offer useful traits, such as abiotic stress tolerance, pathogen
resistance, drought tolerance, and insect resistance. The various
aspects of improved grain composition include increased levels
of specific vitamins (e.g., vitamin A) and minerals (e.g., iron)
and a reduced level of gluten-type protein in barley, permitting
the production of gluten-free beer.
The composition and functional properties of the edible
grains offer a wide diversity of possibilities for new and varied
end uses. Given this genetic diversity, and the consequent
access to the genes responsible, combined with the capabilities
of novel methods of genetic manipulation, we have many
possibilities for extending the compositional diversity of any
grain to provide it with new functional properties.
 THE CEREAL GRAINS | An Overview of the Family of Cereal Grains Prominent in World Agriculture
Figure 17 The inscription on this old plate portrays the ongoing
problem of finding enough grain to provide food for everyone in the
world. The words Altes Brot ist nicht hart kein Brot, das ist hart!
translate as Old bread is not hard no bread, that is hard.
Reproduced from Ng PKW and Wrigley CW (eds.) (2002) Wheat Quality
Elucidation, p. 24. St. Paul, MN: American Association of Cereal
Chemists, with permission.
Exercises for Revision
What features distinguish cereal grains from other food
grains, for example, from the oilseeds and legumes?
What are the distinct meanings of the terms seed, kernel,
corn, and caryopsis?
Which three cereal grains are grown to the greatest extent
worldwide?
Which cereal genus and species is best suited to (a) bread
making and (b) pasta making?
Which cereal grain is favored for making beer; or are several
cereal grains used for this purpose?
How do we come to use the term cereal in common
speech for breakfast foods such as cornflakes and rice
bubbles?
What protein part of wheat flour makes it uniquely suitable
for bread making?
Cereals are members of the flowering plant family. What is
the important function of the cereal flower for our inter-
vention in producing new varieties? Cross-reference to
breeding articles.
Exercises for Readers to Explore the Topic Further
The glutenprotein complex of wheat dough is made up of
many proteins. What are the names and functions of some of
the component proteins? Cross-reference to cereal proteins.
Select several countries and identify their national foods
derived from cereals, especially the species most grown in
each country or region.
85
How are cereal species most similar and most divergent
with respect to the morphology (shape) of the plants, the
grain-bearing head (panicle), and the grain/seed?
Investigate the specific advantages offered by GM cereals.
Which of these are at the research or commercial stages? In
which regions or countries are these permitted?
See also: Carbohydrates: Cereals: Chemistry and Physicochemistry
of Non-starchy Polysaccharides; Food Grains and the Consumer:
Grains and Health; Food Grains: Intolerance, Allergy and
Diseases: Celiac Disease; Non-wheat Foods: Maize: Foods from
Maize; Rice: Chinese Food Uses; Sorghum: Utilization; Processing of
Grains: Cereals: Breakfast Cereals; Extrusion Technologies; Proteins:
The Protein Chemistry of Cereal Grains; The Basics: Grain and Plant
Morphology of Cereals and how characters can be used to identify
varieties; The Grain Crops: An Overview; Grain: Morphology of Internal
Structure; The Grain Chain: The Route from Genes to Grain-Based
Products; The Cereal Grains: African Rice (Oryza glaberrima): A
Brief History and Its Growing Importance in Current Rice Breeding
Efforts; Barley: An Overview of a Versatile Cereal Grain with Many Food
and Feed Uses; Durum Wheat: Overview; Maize: Overview; Millet
Minor: Overview; Millet Pearl: Overview; Oats: Overview; Rice:
Overview; Rye Grain: Its Genetics, Production, and Utilization; Sorghum
Grain, Its Production and Uses: Overview; Teff: Overview; Triticale:
Overview; Wheat: An Overview of the Grain That Provides Our Daily
Bread; Wildrice, Zizania: Overview; Wheat Processing: The Gluten
Proteins of the Wheat Grain in Relation to Flour Quality; Wheat-
Based Foods: Breads; Cakes, Muffins and Bagels; Cookies, Biscuits
and Crackers: Formulation, Processing and Characteristics; Flatbreads
of the World; Noodles: Asian Wheat Flour Noodles; Noodles: Starch;
Pasta; Sour Dough Technology; Tortillas; Cereals: Overview of Uses:
Accent on Wheat Grain.
Further Reading
Abdel-Aal E and Peter Wood P (eds.) (2005) Specialty Grains for Food and Feed.
St Paul, MN: AACC International Press.
Arendt EK and Zannini E (eds.) (2013) Cereal Grains for the Food and Beverage
Industries. Oxford: Woodhead Publishing.
Bhattacharya KR (2011) Rice Quality. Cambridge: Woodhead Publishing.
Champagne ET (ed.) (2004) Rice: Chemistry and Technology, 3rd edn. St Paul, MN:
AACC International Press.
Dendy DAV (ed.) (1994) Sorghum and Millets: Chemistry and Technology. St Paul, MN:
AACC International Press.
Henry RJ and Kettlewell PS (eds.) (1996) Cereal Grain Quality. London, UK: Chapman
and Hall.
Kent NL and Evers AD (eds.) (1994) Technology of Cereals. An Introduction for
Students of Food Science and Agriculture, 4th edn. Oxford, UK: Pergamon Press.
Khan K and Shewry PR (eds.) (2009) Wheat: Chemistry and Technology, 4th edn. St
Paul, MN: AACC International Press.
Shewry PR and Ullrich SE (eds.) (2014) Barley: Chemistry and Technology, 2nd edn. St
Paul, MN: AACC International Press.
Sissons MJ, Abecassis J, Marchylo B, and Carcea M (eds.) (2012) Durum Wheat:
Chemistry and Technology, 2nd edn. St Paul, MN: AACC International Press.
Webster FH and Wood PJ (eds.) (2011) Oats: Chemistry and Technology, 2nd edn. St
Paul, MN: AACC International Press.
White PJ and Johnson LA (eds.) (2003) Corn: Chemistry and Technology, 2nd edn. St
Paul, MN: AACC International Press.
Wrigley CW and Batey IL (eds.) (2010) Cereal Grains: Assessing and Managing Quality.
Oxford: Woodhead Publishing.
 Cereals: Domestication of the Cereal Grains
LA Morrison, Oregon State University, Corvallis, OR, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
As members of the grass family, cereals have inconspicuous
flowers located on a complex inflorescence structure that
can be one of three types: spike, raceme, or panicle.
The florets of cross-pollinating cereals open up at anthesis
to allow the stamens to shed their pollen out into the wind
and air currents, which move the pollen to the stigma of
another floret for fertilization.
Opening of the bracts on grass florets and spikelets during
pollination to both spread and receive pollen fosters the
hybrid speciation that characterizes cereal evolution.
Dispersal strategies in wild progenitors of cereals shed the
mature grain off of the mother plant in a protective dis-
persal unit known as a diaspore, which is formed by the
adherent sterile bracts (glumes, lemmas, and paleas) whose
shape, awns, and bristles aid in its movement to a suitable
habitat for seedling establishment.
Domestication altered the wild dispersal systems of cereal
progenitors by developing forms that retain the grain on the
mother plant while easily releasing the grain during thresh-
ing, the two significant changes that led cereal evolution.
As Old World cereals originating in the Fertile Crescent,
wheat and barley are primary domesticates whereas oats
and rye are secondary forms given their initial status as
weedy wild species infesting early wheat and barley fields.
Maize is the only New World cereal and unusual for its
highly compressed raceme that developed from a pistillate
inflorescence.
Pearl millet and sorghum are drought-tolerant cereals
domesticated in Africa, which spread into South Asia
where they underwent secondary domestication.
Rice is a semiaquatic cereal initially domesticated in south-
ern China, whose phylogeny is characterized by extensive
introgressive hybridization between wild and domesticated
forms and between the two domesticated races indica and
japonica.
Wheat offers the unusual evolutionary example of a multi-
level phylogenetic mosaic built on multiple rounds of
homoploid and polyploid hybrid speciation events.
Weedy cereal relatives have been domesticated as a conse-
quence of human agricultural practices and in some cases
such as pearl millet, have increased their diversity and
adaptability through introgressive hybridization with the
crop to form cropweed races.
While the major cereals represent a large component of
agricultural economies, the minor grains are greater in num-
ber and represent a more diverse distribution, particularly in
subsistence farming systems where they are major food
staples.
Plant breeding applications of the Brachypodium and Setaria
functional grass model systems promise major advance-
ments in improving agronomic and physiological traits of
cereals.
86 Encyclopedia of Food Grains, Second Edition
Realizing the full potential of well-defined phylogenetic
relationships that can guide plant breeders in efforts to
tap the diversity of cereal crop gene pools requires a global
commitment to germplasm conservation.
Learning Objective
To achieve understanding of the mechanisms of domesti-
cation from the standpoint of grass reproductive biology,
the associated natural evolutionary processes such as
hybrid speciation, and the human activities that led the
development of cereal species
Introduction
Cereals comprise a diverse collection of species, all belonging to
the grass family (Poaceae). Collectively, these species cluster in
only four of the 12 grass subfamilies: Chloridoideae, Ehrhartoi-
deae, Panicoideae, and Pooideae. As a group, they are named
for the Roman goddess Ceres who was worshipped as the
mother of agriculture and goddess of grain. The cult surround-
ing Ceres built on her earlier Greek (Demeter) and Egyptian
(Isis) counterparts. In the case of Ceres and Demeter, their
symbols included heads or sheaves of grain, usually wheat for
Ceres and barley for Demeter.
Cereal evolution is a complex subject given the vast array of
grass species adopted into agriculture. Most researchers and
consumers think in terms of the relatively narrow selection of
species that dominate international commerce as dietary sta-
ples and constituents in processed food products. In this select
group are barley, maize, pearl millet, oats, rice, rye, sorghum,
and wheat. Cereals actually comprise a much larger group with
many lesser-known species cultivated and utilized on a local
basis, particularly in subsistence farming cultures.
Inflorescence Structure
Members of the grass family all have inconspicuous flowers
that have been reduced to the essential reproductive structures.
There are no showy bracts, nectaries, and other features to
attract animal pollinators. Unless self-pollinated, cereal species
rely on the wind to spread their pollen. Although grass flowers
all have the same basic design (Figure 1), they can vary in
terms of presence and functionality of the female and male
reproductive structures. Each flower is borne singly in a floral
unit (floret) enclosed by two outer leaf-like bracts (lemma and
palea) (Figure 1). Florets can be bisexual, pistillate (female), or
staminate (male). Fertile florets contain three stamens (filament
and anther) and one pistil (stigma, style, and ovary) with two
tiny, scale-like appendages (lodicules) positioned at its base.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00007-3
 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains 87

Anther
Stigma

Stamen
Pistil
Style
Filament

Ovary
Lodicule
Rachilla

Palea
Lemma

Upper (second)
glume
Lower (first)
glume
Palea

Lemma

Rachilla
joint Callus
Figure 1 Stylized drawings of a grass flower (upper left), floret with adjacent rachilla internode (lower left), and spikelet with subtending rachis
internode (right). Reproduced with permission from Clark LG and Pohl RW (1996) Agnes Chases First Book of Grasses. Smithsonian Press.

The floret is attached to a secondary stem axis, the rachilla. Pollination, Fertilization, and Grain Development
Subtending the rachilla are two empty bracts, the glumes. One or
Cereal species can be either self- or cross-pollinating. In the
more florets, packaged within the glumes, form a spikelet
latter species, the lodicules swell at anthesis, forcing the florets
(Figure 1). Spikelets are alternately positioned at attachment
open to expose the stigmatic surfaces of the pistil to incoming
points (nodes) located along the main inflorescence axis (rachis)
pollen. While the florets are open, the stem-like filaments on
or on branches extending off the rachis. The interval between
the stamens elongate to push the anthers upward and out of
two nodes on the rachis is the internode. Within the spikelets,
the floret where they dehisce and shed pollen into the wind
florets are similarly arranged, alternately along the rachilla.
and air currents. The glumes can bend outward at their bases,
Although grass inflorescence morphology is complex, a sim-
enabling a wider opening of the florets. For self-pollinating
plistic classification designates three basic types (Figure 2),
cereal species whose florets do not stay tightly closed at anthe-
each differing in rachis branch development and spikelet
sis, cross-pollination can occur at low levels. This functional
attachment: (1) panicle with long, open branches or dense
morphology of the floret and spikelet at play during pollina-
branches, each bearing a pediceled spikelet; (2) raceme with
tion sets the stage for the hybrid speciation that characterizes
spikelets borne on short pedicels along the rachis; and (3)
cereal evolution.
spike with spikelets sessile along the rachis.
88 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
(a) (b) (c)
Figure 2 Illustrations of the three general inflorescence types found in
grasses: (a) panicle, (b) raceme, and (c) spike. Reproduced with
permission from Clark LG and Pohl RW (1996) Agnes Chases First Book
of Grasses. Smithsonian Press.
The stigma, a collection of sticky feather-like branches, acts
as the pollen trap (Figure 1). Upon pollination, a pollen tube
grows downward through the stigma and style into the ovary,
which holds a single ovule (egg). Fertilization occurs when the
two pollen sperm nuclei unite, respectively, with the egg and
two female polar nuclei. From this event, a grain develops with
a 2n embryo and 3n endosperm. When fully developed, the
cereal grain is the fruit and seed combined, that is, a specialized
fruit known as a caryopsis, whose outermost layer is formed by
the dried remains of the ovary (pericarp), now adherent to the
seed it surrounds.
Dispersal
Wild relatives of cereal species depend on dispersal strategies to
move the propagule unit enclosing the mature caryopses off
and beyond the mother plant. Propagule units (diaspores) con-
sist of one or more grains surrounded by the dried inflorescence
structures. They are shed from the mother plant by spontaneous
shattering (disarticulation) at abscission zones located at the
nodes along the rachis or rachilla (Figure 3). Disarticulation
is typically described as above (rachilla disarticulates into floret
units) or below the glumes (rachis disarticulates into spikelet,
branch, or inflorescence units). The rachilla or rachis internodal
pieces that remain attached may either subtend or lie adjacent
to the diaspore (Figures 1 and 3). Sterile reproductive units or
nonfloral vegetative structures may also form part of the dia-
spore. Examples include the two sterile awn-like lateral spike-
lets on the wild barley diaspore and the reduced inflorescence
(a) (b) (c)
Figure 3 Disarticulation points along the rachis of a spike inflorescence
type illustrating the location of the break and the internode piece
attached to the diaspore: (a) arrow-shaped diaspore with subtending
rachis piece, (b) cylindrical diaspore with adjacent rachis piece, and (c)
tough rachis with no disarticulation. Disarticulation points along the
rachis for racemes and panicles, as well as along the rachilla for floret
units, are analogous to the patterns illustrated here. Reproduced
with permission from Peterson RF (1965) Wheat: Botany, cultivation,
and utilization. In: Polunin N (ed.) World Crop Books. New York:
Interscience.
branches that form a bristly fascicle (involucre) surrounding
the spikelet on the wild pearl millet diaspore.
Wild species also have developed strategies both to move
diaspores to a favorable habitat and to protect the seed from
predation and environmental damage until seedling establish-
ment. For example, hardened glumes (wild wheat) or a rachis
cupule and glume together (wild maize) can completely enclose
the mature spikelet in a protective package; the lemma and
palea can adhere to the grain (wild barley) or form a hard
shell around it (pearl millet). Diaspore shape and awns, barbs,
and bristly hairs or branches determine the mode of movement
once the diaspore separates from the mother plant. Cylindrical-
shaped (wild wheat: Aegilops tauschii) or angular-shaped (wild
maize) diaspores move by rolling or tumbling across the
ground, floating on water, or getting caught in mud that sticks
to the hides or in the hooves of animals. Arrow-shaped units fall
or are moved point downward into soil cracks (wild rye). Awns,
panicle branches, and bristles subtending spikelets can aid in
the movement of the diaspore to an ideal site for establishment.
 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
Diaspores generally move only a short distance from the mother
plant unless aided by animals, humans, or water.
Awns, which are extensions of the vascular nerves of
glumes and lemmas, can move the dispersal units in a variety
of ways. Natural humiditydrying cycles can cause awns to
spread open and close, thereby moving the diaspore by pas-
sive mechanical force with the seed end pointing forward for
short unidirectional distances or down into the soil (wild
wheat: Triticum urartu). Other hygroscopic awns will coil
and uncoil in response to moisture changes, thereby drilling
the diaspore into the ground (wild oat). Tiny barbs on awns
or bristles can ratchet the movement of diaspores across the
ground for short distances, lock the diaspore in place in the
soil, or catch the diaspore on the fur of animals or clothing of
humans. Awns can also vibrate in the wind to push the
diaspore downward into the soil or soil crack. Lightweight,
feathery bristles of a fascicle can move a diaspore in the wind
(wild pearl millet).
In contrast to their wild relatives, domesticated cereal species
retain the mature caryopsis on the plant and have more easily
extractable grains due to reduced or absent lemmas, paleas, and
glumes. Thus, humans must remove the mature grains from the
mother plant by hand or mechanical harvesting, thresh the
harvested heads to free the grain from their enclosing bracts,
protect the grain from predation and environmental threats,
and plant it for a new crop. Some of the more primitive cereal
species still retain wild-type features. For example, semibrittle
forms of emmer and spelt wheat (T. dicoccum and T. spelta) can
naturally disarticulate or break under relatively little mechanical
pressure into spikelet diaspores enclosed by tough glumes.
Evolution and Domestication
Domestication constitutes an evolutionary change and specia-
tion mediated by humans acting as selection agents. For the
cereal grains, early humans channeled the inherent genetic
variation of wild grasses in the direction of agricultural traits
that eased work to cultivate, harvest, and extricate grain. Then,
through their ancient migrations and trade routes, they spread
domesticated cereals and associated weed species beyond the
original sites of domestication. With the expansion of early
agriculture, opportunities opened for hybrid speciation events
between otherwise allopatric species or between weedy wild
forms and crop species. These expanded associations led to
new domesticated forms (for example, T. aestivum) or intro-
gressed forms with more complex phylogenies (maize, rice, and
sorghum). Ancient farmers further exploited the domestication
process by developing unique land races adapted to the grow-
ing conditions and food needs of their locales. As a result of
these factors, discerning phylogenetic relationships can be chal-
lenging. Thus, taxonomic classifications constructed to reflect
phylogenetic evidence can be dynamic, changing as the evi-
dence changes. The practice of classifying the wild and domes-
ticated cereal forms as one biological species, usually named
with the domesticated species name, can lead to confusion if
the wild versus domesticated forms at the infraspecific levels of
subspecies or variety are not specified (for example, T. turgidum
and Sorghum bicolor can refer broadly to both wild and domes-
ticated taxa).
89
Domestication Syndrome
Under domestication, features of dispersal, reproduction, plant
growth, and survival of wild grasses were transformed into
desirable agricultural traits designated as the domestication syn-
drome. The two leading transitions were: (1) the retention of
the mature grain on the mother plant due to the development
of a nonshattering, tough rachis and (2) the easy release of the
grain from the bracts due to morphological changes reducing
or eliminating glume tenacity and adherence of the lemma and
palea to the mature grain. Following these initial changes,
humans manipulated a suite of desirable physiological and
agronomic traits to further advance early forms into fully agri-
cultural species:
(1) Reduction of seed dormancy either by loss of physiological
dormancy or by loss or reduction of enclosing bracts that
contained germination inhibitors
(2) Increase in seed production by development of reproduc-
tively functional florets from sterile or vestigial ones
(3) Increase in seed size
(4) Synchronous development of tillers and ripening of the
grain
(5) Changes in photoperiod sensitivity
(6) Greater agricultural fitness for soil and climatic conditions
and pest resistance.
The speed and spread of the transition to agriculture is still
debated with differing interpretations of domestication as
rapid versus gradual and spread of initial domesticates as an
outward diffusion from a primary place of origin versus a series
of relatively concurrent events in different locales.
Reduction or full loss of the rachis nodal abscission zones is
responsible for the tough rachis trait. Semishattering forms
naturally break into diaspores in the upper portion of the
inflorescence or break when pressure is applied in partially
formed abscission zones. Changes in bract morphology led to
free-threshing cereals. The tough tenacious glumes softened or
became more brittle, losing their clasping feature that enclosed
the spikelet at maturity. Lemmas and paleas lost their tendency
to adhere to the mature grain. Size reduction, anatomical thin-
ning, and increased brittleness of the bracts allowed them to
easily fall away with mechanical threshing to release a naked
grain. The term hulled cereal refers to different protective features
of the bracts that have not completely lost wild-type features:
Hulled grains either are encased within the lemma and palea,
which either form a hard shell around the grain (pearl millet) or
adhere to the grain surface (barley); hulled spikelets have hard-
ened, tough glumes that tightly enclose one or more grains in a
spikelet package (spelt wheat, that is, T. spelta).
Domestication Parallels: Crop and Weed Races
Domestication followed different routes for each cereal species
and did not necessarily involve a straightforward, one-time
transition from wild to domesticated species. Introgressive
hybridization has played a significant role and followed a
variety of pathways. Domesticated races intercrossed with
each other (rice), with weedy wild relatives (barley), or with
wild species related to the original wild progenitor (sorghum,
pearl millet, and maize). Polyploid speciation in the wheats
90 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
and oats illustrates how new genome material introduced
through interspecific hybridization formed new species. These
types of events increased the narrowed diversity arising from
the domestication bottleneck, thereby expanding the adaptabil-
ity of domesticated forms and their distribution ranges.
The common element among wild cereal progenitors is
their amenability to cultivation in human-modified habitats.
Along with domesticated forms came weeds, the unwanted
species that also thrive in these habitats. The development of
weed races parallels crop evolution from the standpoint of
unintended human selection of weedy forms through agricul-
tural practices and introgressive hybridization between the
crop species and its weedy wild relatives. Weed races in crop
weed complexes can be serious pests of agricultural fields as is
the case for sorghum and rice. Or, weeds can develop into crop
species as happened to the weedy races of oats and rye that
infested early wheat fields. As wheat and barley agriculture
spread outward from Southwest Asia, wild oat and rye traveled
along as grain contaminants into regions where they developed
into domesticated forms suited to growing conditions that
were less favorable to wheat or barley.
Major Cereal Species
Major cereal species (Table 1) are those that are in interna-
tional commerce and serve as dietary staples beyond a local-
ized region. Wheat and barley are temperate cereal crops of the
Fertile Crescent, and along with oats and rye, formed the major
grain complex for Western Asia, North Africa, and Europe.
Maize and rice are the tropical cereals upon which civilizations
in the New World and Eastern Asia developed. Although sor-
ghum, pearl millet, and African rice, all tropical African species,
have historically been given a lesser role, they are equally
important in the establishment of early human civilizations
in Africa and South Asia.
Barley
Domesticated barley (Hordeum vulgare) is an annual diploid
species derived from the wild progenitor species
H. spontaneum. In preagricultural archaeological finds, barley
is a universal companion of wheat. While considered inferior as
Table 1 Major cereals and their wild progenitors
Domesticated
Common name species
Barley Hordeum vulgare
Common oat Avena sativa
Maize Zea mays
Pearl millet Pennisetum
glaucum
Rice Oryza sativa
Cereal rye Secale cereale
Sorghum Sorghum bicolor
Durum wheat Triticum durum
Bread wheat Triticum aestivum
a
Approximate dates of first appearance of domesticated cereals based on current reports of archeological and genetic evidence.
a staple, barleys greater adaptability to poor soils and tolerance
of soil salinity increased its culture in areas and climatic condi-
tions unfavorable to wheat. The barley spike (or spike-like
raceme) has three spikelets at each rachis node. Spikelets con-
tain one floret. Their row-like appearance along both sides of
the spike and floret fertility is the basis for dividing domesti-
cated barley into two groups: (1) two-rowed barley with a
central fertile spikelet and two adjacent sterile spikelets, each
containing a male floret (primitive form; subsp. distichum)
and (2) six-rowed barley with all three spikelets fertile
(advanced form; subsp. hexastichum). Most domesticated bar-
ley has a hulled grain (adherent lemma and palea), although
there are naked six-rowed cultivars with the botanical ranking
of var. coeleste (var. nudum).
The domestication of barley has been traditionally treated
as a single event occurring in the Fertile Crescent. Although
current genetic evidence has supported one independent ori-
gin, recent studies show a second primary center of origin
located in the region east of the Zagros Mountains. Thus, the
barleys of Europe and North Africa developed from the Fertile
Crescent line whereas those of Central and East Asia developed
from the Zagros Mountains line. The H. spontaneum wild weedy
complex spread from its primary habitats in the Fertile Cres-
cent to secondary habitats westward in the Mediterranean
Aegean regions and eastward across Central Asia to Tibet.
The weedy forms (H. spontaneum) have played a role in
building diversity through introgressive hybridization with
domesticated barley. For example, the fragile-rachis, six-
rowed domesticated barley discovered in Tibet, and originally
proposed as a distinct wild progenitor species (H. agriocrithon),
is an introgressed form. Although barley has a lengthy agricul-
tural history and development of an extensive number of
cultivars and land races, its morphological diversity is relatively
limited.
Maize
Among the domesticated cereals, maize (Zea mays subsp. mays)
has an unusual reproductive biology. It is a diploid, cross-
pollinating monoecious species (staminate and pistillate inflo-
rescences on the same plant) whose pistillate inflorescence is a
highly compressed raceme that develops into a cob bearing
naked grains. Maize originated from a single domestication
Region of primary Date of primary
Wild progenitor species domestication domesticationa
Hordeum spontaneum Southwest Asia 10 2009550 BP
Avena sterilis Europe 76007400 BP
Zea mays subsp. parviglumis Mesoamerica 8700 BP
Pennisetum violaceum West and Central Africa 4800 BP
(P. mollissimum)
Oryza rufipogon Southeast Asia 800013500 BP
Secale vavilovii Southwest Asia 94508450 BP
Sorghum arundinaceum Northeast Africa 8000 bp
Triticum dicoccoides Southwest Asia 10 2009550 BP
unknown West Asia 80007000 BP
 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
event in the lowland regions of southwestern Mexico where its
wild annual progenitor, Balsas teosinte (Z. mays subsp. parvi-
glumis) is still endemic. Interestingly, maize ancestry has
proved to be more complex than a straight wild to crop species
development. Genetically, all maize cultivars in the Americas
can be traced to highland maize, which also is the maize most
closely related to the wild progenitor parviglumis. This ancestral
relationship sets up a geographic contradiction due to a low-
land origin but a highland ancestral line. With parviglumis
distribution restricted to mid- and lowland regions, a highland
ancestral form is difficult to explain. Recent genetic research,
which tracks with ecological and archaeological findings, has
helped resolve the conflicting evidence by identifying a sec-
ondary teosinte progenitor, Z. mays subsp. mexicana. Highland
teosinte grows sympatrically with highland maize and is inter-
fertile with maize and parviglumis. While mexicana is not the
direct progenitor of maize, evidence now points to gene flow
from this wild relative as the explanation for highland maize
holding the ancestral genetic position. In this respect, maize
offers an informative example of the complexities of discerning
crop origins when geography and introgressive hybridization
between the domesticated form and secondary wild relatives
play a role.
Pearl Millet
Millet in a broad sense comprises a loose grouping of small-
seeded, drought-tolerant species that can be cultivated on poor
soils (Figure 4). Collectively, the millets belong to nine genera
in the grass subfamilies Paniceae (eight genera) and Chloridoi-
deae (one genus) (Table 2). Due to their small size and a
hardened lemma and palea, millet grains are insect-resistant,
a feature that adds to their value for the African and Asian
cultures that cultivate them. Only pearl millet (Pennisetum
glaucum) is a major cereal grain on an international scale. It is
a largely cross-pollinating plant due to the unsynchronized
maturity of the respective female and male florets. A diploid
Figure 4 Diversity and form in the millets. From left to right: panicles of
pearl millet, broomcorn millet, foxtail millet, and Japanese millet.
Reproduced with permission from Hancock JF (2004) Plant Evolution
and the Origin of Crop Species 2nd edn., Portland, OR: Book News Inc.
91
annual, pearl millet was domesticated in the arid regions of
West Africa approximately 5500 BP, moving to East Africa and
then India 3000 BP and to Southern Africa 2000 BP. In West
Africa, pearl millet still forms an integral component of decrue
(flood stage to dryland) agriculture in the Central African river
deltas where it is cultivated along with African rice (Oryza
glaberrima) and sorghum (Sorghum bicolor). Pearl millet is
more drought-tolerant than sorghum and maize.
Low genetic variation suggests a relatively limited region of
origin, although there are a diversity of morphological forms
varying in seed shape, seed color ranging from red to gray to
black, and a spike-like compressed panicle that can measure up
to 2 m. Pearl millets are grouped into four ecogeographic races
with characteristic morphological traits: typhoides (Africa and
India), nigritarum (eastern Sahel), globosum (western Sahel),
and leonis (West African coast). Active introgression makes
it difficult to draw clear phylogenetic lines between pearl
millet, its wild progenitor P. violaceum, and the weedy species
P. sieberanum, which evolved from crosses between P. violaceum
and domesticated millet. Crop mimicry is a problem in African
fields of pearl millet where weed races of P. sieberanum, known
as shibras, are almost indistinguishable from the domesticated
species.
Oats
Oats form a polyploid series (2, 4, and 6) of self-
pollinating species following different evolutionary lines. Four
genomes characterize this group: A and C evolved independently
of one another; B and D are derivatives of A. The dominant
domesticated species is the common oat, Avena sativa (ACD),
which developed from a Southwest Asian ecogeographic race of
the wild progenitor species A. sterilis. The less important but
closely related red oat, A. byzantina ( A. sativa subsp. byzantina)
was likely derived from a different disjunct regional A. sterilis
race. The diploid sand oat, A. strigosa (As, a variant of A), evolved
in Spain from the wild form of this species. The semi-
domesticated, weedy tetraploid Ethiopian oat, A. abyssinica
(AB), evolved in Ethiopia from the wild species A. vaviloviana.
A. strigosa and A. abyssinica did not play a role in the evolution of
the hexaploids.
More water-tolerant than the other major temperate cereals,
common oat is suited to cultivation in temperate regions with
wet, cool summers. It has an open, nonshattering panicle with
hulled (adherent lemma and palea) seeds except for two naked
forms: the hexaploid A. sativa subsp. nuda with origins in
Mongolia and northwest China and the diploid A. strigosa
subsp. nudibrevis, which developed in England.
With its center of origin located in the western Mediterra-
nean, A. sterilis has a distribution extending from the Atlantic
coast to the Zagros Mountains. A species of primary habitats
often growing alongside wild wheat and barley, wild oat also
populates abandoned and actively cultivated agricultural land
and sites associated with human disturbance. Its initial domes-
tication began as a weed in the newly opened habitats of
cultivated wheat and barley fields. As A. sterilis spread eastward
and westward with wheat and barley agriculture, a complex of
weed races evolved alongside the domesticated common oat,
which began to appear c. 40003000 BP in Europe. In addition
to these domesticated and weedy A. sativa forms, the A. sterilis
92 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains

Table 2 Minor cereal grains either cultivated or harvested as cultivated or uncultivated wild species

Cereal Species Region where cultivated or collected Domestication statusa

Adlay millet Coix lacryma-jobi South Asia, East Asia Domesticated

Fonio Digitaria
Raishan D. cruciata South Asia, East Asia Domesticated
White Fonio D. exilis West Africa Domesticated
Black fonio D. iburua West Africa Domesticated
Polish millet D. sanguinalis Europe, Asia Domesticated
Wild fonio D. fuscescens West Africa Wild
D. longiflora
Millets
Guinea millet Brachiaria deflexa West Africa Domesticated
Browntop millet Brachiaria ramosa South Asia Domesticated
Sawa millet Echinochloa frumentacea Asia Domesticated
Shama millet Echinochloa colona Africa Wild
Japanese millet Echinochloa esculenta East Asia Domesticated
Finger millet Eleusine coracanab Africa, Asia Domesticated
Desert panic Panicum laetum Africa Wild
Broomcorn millet Panicum miliaceum Asia Domesticated
Mexican panicgrass Panicum hirticaule North America Domesticated
Little millet Panicum sumatrense Asia Domesticated
Kodo millet Paspalum scrobiculatum South Asia Domesticated
Kodo millet Paspalum scrobiculatum Africa Semidomesticated
Foxtail millet Setaria italica Southeast Asia Domesticated
Oats Avena
Abyssinian oat A. abyssinica East Africa Semidomesticated
Red oat A. byzantina Mediterranean Basin, West Asia Domesticated
Sand oat A. strigosa Southern Europe Domesticated
Rice
Wild rice Oryza barthii West Africa Wild
Wild rice Oryza punctata East Africa Wild
African rice Oryza glaberrima West Africa Domesticated
Southern wild rice Zizania aquatica North America Wild
Northern wild rice Zizania palustris North America Wild, domesticated
Rye Secale segetale West Asia Semidomesticated
Sand Bur Cenchrus biflorus Africa, Asia Wild
Tef Eragrostis tef East Africa Domesticated
Triticale Triticosecale Worldwide Domesticated
Wheat (2) Triticum
Small spelt T. monococcum Eurasia Domesticated
Wheat (4) Triticum
Emmer T. dicoccum Eurasia Domesticated
Isfahan wheat T. ispahanicum South Asia Domesticated
Georgian emmer T. paleocolchicum Western Asia Domesticated
Ethiopian wheat T. aethiopicum East Africa Domesticated
Khorasan wheat T. turanicum South Asia Domesticated
Persian wheat (4, 6) T. carthlicum Southwest Asia Domesticated
Polish wheat T. polonicum Southwest Asia, Mediterranean Basin Domesticated
Rivet wheat T. turgidum Eurasia Domesticated
Timopheevs wheat T. timopheevii West Asia Domesticated
Wheat (6) Triticum
Spelt wheat T. spelta Eurasia Domesticated
Macha wheat T. macha West Asia Domesticated
Vavilovs wheat T. vavilovii West Asia Domesticated
Shot wheat T. sphaerococcum South Asia Domesticated
Zhukovskys wheat T. zhukovskyi West Asia Domesticated
a
For additional information, see Luven P (1988) Traditional Food Plants. FAO Food and Nutrition Paper 42. FAO; Vietmeyer (Director) ND (1996) Lost Crops of Africa: Grains, vol. 1.
Washington, DC: Board on Science and Technology for International Development, National Research Council, National Academy Press; Wiersema JH and Leon B (2013) World
Economic Plants: A Standard Reference, 2nd edn. New York: CRC Press, Taylor & Francis Group. Williams JT (ed.) (1995) Cereals and Pseudocereals. London: Chapman and Hall,
and website sources listed at the end of this article.
b
Member of the subfamily Chloridoideae. All other millet species belong to the subfamily Paniceae.
 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
line gave rise to other interfertile weedy wild hexaploid species:
A. ludoviciana, A. occidentalis, and A. fatua. Of these, A. fatua is
an aggressive agricultural weed whose extensive distribution is
aided by its spring type habit, that is, it does not require
vernalization. Similarly, the wild tetraploid A. barbata (AB) is
a noxious weed worldwide wherever it has been introduced
into Mediterranean-like habitats.
Rice
Although there are two species of rice, only one of these, Oryza
sativa, which was domesticated in Asia, has become a major
cereal staple in international agriculture. In contrast,
O. glaberrima is a minor cereal whose cultivation is confined
to a small area in the equatorial belt of its native West Africa.
Both domesticated species and their wild relatives are diploid,
semiaquatic plants whose premodern cultivation was tied to
seasonal flooding and rains. Of the two, O. sativa has a greater
climatic and latitudinal diversity with a range of forms found
from tropical to temperate zones and cultivated in different
habitats defined by water availability: semiaquatic, either
rainfed or irrigated (lowland); deep water (floating); coastal
wetland; and dryland (upland). It is a highly diverse species
with two major races, sometimes treated as subspecies indica
and japonica. Based on current archeological and genetic evi-
dence, the japonica form was first domesticated from its wild
relative O. rufipogon in the middle area of the Pearl River in
southern China c. 13 5008000 BP. Indica subsequently devel-
oped from hybridization events between O. rufipogon and the
initial japonica cultivars as they spread into Southeast and
South Asia. Diversity within the two races and distinctive dif-
ferences between them are attributable to introgression
between the wild and domesticated forms as well as introgres-
sion of indica into japonica forms. This variation is grouped into
five subpopulations whose boundaries are not always well
defined: temperate japonica, tropical japonica (javanica), bas-
mati-type (aromatic), ausboro, and indica.
Domestication of the considerably less diverse O. glaberrima
is dated to 3500 BP in the upper Niger River Delta. Its direct
wild progenitor is the wild annual species, O. barthii, which
populates savanna water holes and has weedy forms that infest
rice fields located in the high rainfall belt of the African
savanna. Oryza sativa was introduced into West Africa from
Europe in the fifteenth century CE and now predominates.
Hybrid cultivars known as NERICA (NEw RICe for Africa)
developed from crosses combining the stress and disease tol-
erance of O. glaberrima with the yield potential of O. sativa
show promise for improving food security in the rice-growing
areas of Africa.
Rye
Cereal rye (Secale cereale) belongs to a small Old World genus
of mostly annual diploid species whose phylogenetic relation-
ships are complicated by nondomesticated taxa with an obli-
gate cross-pollinating reproductive biology and perennial
habit. Its spikes are nonshattering, producing free-threshing
grains at maturity. Cultivation extends into areas requiring
winter hardiness, drought-resistance, and tolerance of sandy
and acid soils. Preagricultural forms of wild-type rye found in
93
Neolithic archaeological sites suggest its status as a tolerated
weed of wheat fields. Domesticated rye began to appear circa
95008500 BP and likely originated in eastern Turkey and
western Armenia. From there, it spread across the Mediterra-
nean Basin, into Europe, and eastward into Central Asia. An
interfertile crop complex of wild, weedy, and domesticated
races, cereal rye is classified into four groups according to
spike fragility or shattering: (1) nonshattering domesticated
varieties (S. cereale subsp. cereale); (2) weedy forms (segetale)
that mimic wheat in spike and grain morphology and are
harvested and threshed as part of the wheat crop; (3) semi-
shattering weeds (afghanicum) whose upper spike shatters,
spreading seed in the field while the lower, intact portion is
harvested with the wheat; (4) shattering wild-types (ancestrale)
found in secondary habitats whose spikes fully disarticulate
into spikelet dispersal units enclosed by brittle glumes.
Included in the wild type group is the wild species S. vavilovii
(S. cereale subsp. vavilovii) that grows in primary basaltic
bedrock habitats in Turkey. Evidence points to S. vavilovii as
ryes wild progenitor.
The wild annual relatives S. iranicum and S. sylvestre, both
self-pollinating species, are reproductively isolated from cereal
rye. All the wild perennial rye forms belong to S. montanum
(S. strictum), a diverse species with intergrading and interfer-
tile ecogeographic races, among them dalmaticum (Mediterra-
nean Basin), anatolicum and ciliatoglume (Anatolia), and
kupriyanowii (Iran and Transcaucasia). On the Anatolian
plateau of Turkey where weedy cereal rye and S. montanum
come into contact, the two species hybridize to produce inter-
mediate, introgressed forms.
Sorghum
Sorghum (S. bicolor) is a drought-tolerant, self-pollinating C4
cereal grown throughout the arid and semiarid tropics and
particularly important in subsistence farming cultures. In cul-
tivation, sorghum is treated as an annual although most forms
are perennial. Domestication took place in the northeastern
savanna/steppe region of Africa in an area stretching from
present-day Sudan to Ethiopia around 8000 BP. From there,
sorghum spread to West Africa, down into southern Africa, and
eastward to India, Pakistan, and China. The early extensive
range of sorghum has led to a diverse mix of regional forms.
The wild progenitor S. arundinaceum; (S. bicolor subsp. verti-
cilliflorum), is still found in primary grassland habitats in East
Africa. Secondary introgression from hybridization events
between domesticated sorghum and S. arundinaceum may be
responsible for expanding sorghums ecological range from
arid savanna habitats into areas with higher rainfall and
humidity where it can tolerate up to 5000 mm of rain during
the growing season. The cropweed complexes of sorghums
native Africa also characterize its cultivation in Asia, Australia,
and the Americas due to the introduction of contaminated seed
stocks. Thus, the bidirectional gene flow between the crop and
weed essentially occurs worldwide.
Through domestication, the shattering, open panicles of
S. arundinaceum evolved toward nonshattering, compact pani-
cles with larger grains. Traditionally, sorghum is classified into
five races bicolor, guinea, caudatum, kafir, and durra and ten
two-by-two race combinations, all 15 with a distinctive
94 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
Bicolor
Caudatum Kaffir
Figure 5 Panicles and mature grains still enclosed by their glumes for each of the five sorghum races. Reproduced with permission from Luven
P (1988) Traditional Food Plants. FAO Food and Nutrition Paper 42. FAO.
ecogeographic range and place in indigenous African and Asian
agriculture as well as morphological and use differences
(Figure 5). Bicolor, the most primitive and similar in appear-
ance to wild sorghum with its loose, open panicle, was the first
domesticated race from which the other three races were
derived. Caudatum, which is found from Lake Chad to Ethiopia
in bicolors primary domestication zone, is unique for its grain
shape flat on one side, convex on the other. Kafir is associated
with the Bantu-speaking peoples of East and Southeastern
Africa who migrated there from West Africa. Durra, which has
dense, compact head-like panicles, was developed in India,
where it is now largely grown. African durra likely arrived
from India during Islamic migrations into Africa. Recent
research confirms that the fifth race (guinea), now designated
the guinea margaritiferum group, is the product of a second
domestication event in West Africa. Divergent from the four
other sorghum races, the guinea margaritiferum group is inter-
mediate between sorghum and its wild progenitor and suited to
the poor soils and erratic rainfall conditions of West Africa.
Guinea margaritiferum sorghum and the wild Southeast Asian
S.R.
Guinea
S.R.
Durra
species S. propinquum, which is interfertile with domesticated
sorghum, both offer new, untapped genetic resources for
breeding programs.
Wheat
The wheats comprise a large allopolyploid complex of wild and
domesticated species taxonomically classified in the genera
Triticum (wild and domesticated) and Aegilops (wild). They
are annual, self-pollinating species. Together, hexaploid bread
wheat (T. aestivum) and durum wheat (T. durum; T. turgidum
subsp. durum) are the two major wheats cultivated worldwide,
ranking third in cereal production after corn and rice. The
evolution and domestication of wheat traces an interesting
path in diploid genome development and polyploid specia-
tion. Bread wheats three ancestral genomes A, B, and D are
closely related. A and B diverged approximately 7 million years
ago (mya), giving rise to the D genome 56 mya through
homoploid speciation. The wild diploid species that carry the
respective genomes are T. urartu (A), Ae. speltoides, the putative
 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
B donor species whose subgenome within the Aegilops genus is
traditionally distinguished as S: and Ae. tauschii (D). The evo-
lutionary lines leading from ancient ancestral diploid wheats
to tetraploid and hexaploid wheats constitute a multilevel
phylogenetic mosaic built on multiple rounds of homoploid
and polyploid hybrid speciation events.
The diploid progenitor T. urartu has no domesticated form.
T. dicoccoides (T. turgidum subsp. dicocccoides), the wild tetra-
ploid wheat (BA), originated from an Ae. speltoides x T. urartu
cross. Emmer wheat (T. dicoccum; T. turgidum subsp. dicoc-
cum) (Figure 6) was domesticated from T. dicoccoides circa
10 000 BP in the Fertile Crescent. A primitive form, emmer
has a semifragile spike and hulled grains (clasping glumes).
Its spread eastward brought it into contact with Ae. tauschii,
which populated primary habitats in the Caspian Sea Belt. The
hexaploid hybrid speciation event that formed T. aestivum
(BAD) (Figure 6) likely took place along the edges of cultivated
fields where emmer and weedy Ae. tauschii subsp. strangulata
came into contact. Bread wheat has no known wild hexaploid
progenitor. Evidence suggests that the preliminary hexaploid
form was a primitive hulled T. spelta type. Naked T. aestivum
forms likely developed within a short period afterwards
c.80007000 BP. European forms of T. spelta may have arisen
independently from T. aestivum  T. dicoccum crosses.
The diploid and tetraploid wheats of the bread wheat line
are suited to the Mediterranean-type habitats of Southwest Asia
with mild winters and dry summers. The addition of the D
genome to hexaploid forms extended the ecogeographic range
Figure 6 Diversity within the wheats. From left to right:
T. monococcum, T. dicoccum, and T. spelta (all hulled wheats),
T. polonicum (Polish wheat), T. turgidum (rivet wheat: branched form),
T. aestivum (compactum form), T. durum, T. aestivum (awned*:
Turkey), and T. aestivum (awnless: Wilhelmina). (Adapted from The
Field Museum, negative B79676.) *Awns on Turkey are barely visible
due to photographic quality and their smaller diameter and weaker
anatomy relative to the awns on the T. durum specimen to its left.
95
to continental temperate regions with colder winters and
humid summers. As wheat cultivation of the respective tetra-
ploid and hexaploid forms diffused across Asia, Europe, and
North Africa, humans developed more advanced, specialized
naked grain forms (for example, tetraploid durum and rivet
wheats) (Figure 6; Table 2) as well as a diverse array of land-
races unique for their regional distribution and morphological
and agronomic features. Of note, the transition from wild-type
shattering spikes and hulled grain to tough rachis, free-
threshing domesticated forms follows two paths in the bread
wheat line. The spikelet dispersal units for T. urartu and
T. dicocccoides are wedge-shaped, with the rachis internode
piece subtending the spikelet. In the case of the semishattering
T. spelta, its hulled spikelet units are cylindrical (barrel-
shaped), with the rachis internode adjacent to the spikelet
(Figure 3). Aegilops tauschii also exhibits the cylindrical mode
of spikelet dispersal. All other Triticum (sensu stricto) wild
wheats shatter into the wedge units.
Two other evolutionary lines characterize the wheats. A
different hybrid speciation event between T. urartu and Ae.
speltoides produced the wild tetraploid species T. araraticum
(GA). While T. araraticum is almost identical in morphology
to T. dicoccoides and shares the same distribution ranges
in Turkey, Iraq, and Iran, the two species are reproductively
isolated from each other. Thus, the B and G assignments for
the Ae. speltoides genome contribution recognize the respective
origins for the two tetraploid species. Two domesticated forms
complete this evolutionary line: T. timopheevii (GA) and
T. zhukovskyi (GAAm). Triticum zhukovskyi has no known wild
hexaploid progenitor. The third evolutionary wheat line has
two diploid species: the wild T. boeoticum and its domesticated
offshoot, T. monococcum (Figure 6). Their A genome is different
from that found in the bread wheat line and variously noted at
Ab or Am. Triticum monococcum (einkorn) is an important player
in the early archaeological record but today is considered a relic
domesticated species.
Minor Cereals
The major cereals discussed earlier in the text represent a small
sampling of the diverse collection of grass species that have
been domesticated. Table 2 provides a partial listing of lesser-
known cereals that are currently, or at one time were, impor-
tant crops or are still harvested from wild stands. Some of the
domesticated cereals are cultivated on an extensive scale or
across large regions (foxtail millet). Others are locally
restricted: white fonio in West Africa (Figure 7) and raishan
in India. Their importance is measured by their role as dietary
staples for the peoples who cultivate them. Many of these
species have been supplanted by the major cereals. For exam-
ple, broomcorn millet (Panicum miliaceum), once cultivated
widely in Europe and the United States, has declined in acreage
and use for human consumption. As discussed in the preceding
text, African rice (O. glaberrima), an indigenous species of
decrue agriculture on the river deltas of Central Africa, is
being replaced by O. sativa and NERICA hybrids. On the
other hand, some minor grains are popular in Western spe-
cialty markets, for example, tef and the hulled wheats farro
(T. dicoccum) and spelt (T. spelta).
96 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
Figure 7 Vegetative and reproductive structures of white fonio
(D. exilis). The inflorescence is a digitate raceme with a winged rachis.
Reproduced with permission from Haq N (1995) A neglected cereal and
two promising ones. In: Willams JT (ed.) Cereals and Pseudocereals.
Chapman and Hall.
Wild grasses and partially domesticated grasses still play
an important role in the diet for subsistence cultures. West
Africans gather the wild fonio species D. fuscescens and
D. longiflora in famine times and benefit from harvestable
semidomesticated kodo millet infesting their fields if they
lose the rice crop. The weedy oat species, A. abyssinica, forms
an integral component of the grain harvested from wheat and
barley fields in Ethiopia. Subsistence farmers on the Anatolian
plateau of Turkey allow weedy forms of S. cereale, which are
more cold- and drought-tolerant than the wheat crop that they
infest. In bad years, this harvestable weed rye crop provides the
wheat of Allah for this traditional culture.
Research Trends
Fully characterizing the genetic diversity of cereal species and
that of their wild and weedy relatives is a primary focus of
current cereal research. Along with genome sequencing
projects that are underway or completed, the development of
the annual diploid Brachypodium distachyon as a model grass
system will open avenues for discerning genetic pathways and
gene functions for crop traits such as plantpathogen interac-
tions, yield, and stress tolerance. Where the Brachypodium
model will serve for photosynthetic studies of C3 cereals (rice,
barley, wheat, rye, and oats), the C4 model species Setaria
viridis will do the same for maize, millet, and sorghum. Both
grass models will provide the functional framework for
moving the more efficient C4 system into wheat, rice, and
barley and in developing the promising C4 energy crops such
as switchgrass (Panicum virgatum) and giant miscanthus
(Miscanthus  giganteus).
Phylogenetic research is at various stages of progress depen-
dent on the complexity of the cereal genomes. Availability of
research funding is also a factor governed by position in inter-
national cereal markets. For example, pearl millet is lower on
the funding scale than corn, rice, and wheat. With robust
phylogenetic trees tracing a well-defined picture of relation-
ships, cereal breeders will have a more reliable roadmap for
efficiently exploiting genetic diversity to improve crop traits
and create new cereal crops. The combined potential of the
BrachypodiumSetaria grass model systems and phylogenetic
studies promises a future of expanded efforts in high-tech
cereal breeding, which will become increasingly more impor-
tant for addressing the food security issues of growing popula-
tions, changing climatic conditions, eroding aquifers, and loss
of arable land. An underlying factor in meeting these chal-
lenges will be the availability of germplasm resources. Success-
fully advancing the possibilities of modern domestication
requires an active effort to preserve the existing crop wild
species diversity. Historically, germplasm conservation has suf-
fered from a lack of consistent funding and a coordinated
global commitment. Realizing the full potential of twenty-
first century cereal breeding will require a top priority status
for germplasm conservation programs equal to todays inter-
national research collaborations driving genome mapping
initiatives and grass model systems development.
Exercises for Revision
How does the functional morphology of grass flowers influ-
ence the natural speciation events that shaped the phylog-
eny of wild and domesticated grain species?
Using the example of maize, sorghum, and wheat, explain
the morphological changes under domestication that altered
their respective inflorescences from the wild-type raceme,
panicle, and spike to the advanced domesticated forms.
What is the significance of the development of grain reten-
tion and easily threshed grains in the early development of
agriculture and why did these domestication syndrome
traits predate changes in dormancy, seed size, etc.?
Take one of the positions in the debate over the speed and
spread of domestication and explain why it is the likely
scenario in the development of agriculture.
What is a weed race and what kinds of roles do weeds play
in domestication and the development of agriculture?
How has introgressive hybridization played a role in cereal
evolution and what are examples of the different mechanisms
by which introgression has influenced the phylogenetic rela-
tionships in wild and domesticated crop complexes?
How did their respective evolutionary pathways differ for
the Old World Founder crops barley and wheat?
Explain the features of maize domestication that distin-
guish it from the other major cereals and clarify the
 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
differences between the human-mediated changes and
those that would have happened by natural biological
processes.
How did phenotypic diversity in pearl millet and sorghum
become extensive and what features of their domestication
pathways have encouraged these trends?
How did habitat and the geographic spread of early domes-
ticated forms affect rice diversity and its phylogenetic
relationships?
Although African rice is a minor grain, why is it important
and what are the negative effects of its reduction in acreage
in Africa?
Why are minor grains greater in number and what is their
importance to the regions where they are cultivated?
How will genetic evidence of phylogenetic relationships
advance development of cereal cultivars that will improve
food security under changing climatic conditions and loss
of arable land?
Why is preservation of germplasm diversity, particularly
wild cereal relatives and land race cultivars, of critical impor-
tance for plant breeding and what role can molecular genet-
ics play in building a more complete picture of cereal
diversity?
How will a functional grass model guide future cereal
research to improve agronomic and physiological traits
and what benefits will be gained from moving a C4 photo-
synthetic system into rice and wheat?
Exercises for Readers to Explore the Topic Further
Examine the advantages of multidisciplinary arch-
aeologicalgenetic studies in discerning the timing of
domestication and the location of primary and secondary
centers of cereal domestication and discuss how this work
will inform germplasm conservation strategies and plant
breeding programs.
NERICA hybrids combine the yield benefits of Asian rice
with the drought tolerance and weed and pest resistance of
African rice. Consider the advantages and disadvantages of
NERICA introductions relative to a balance between diver-
sity conservation and food security in Africa.
Explore the role of mapping initiatives and other compre-
hensive genetic studies of cereal species in advancing mod-
ern agriculture. What is the importance of educating the
public in their importance to global food security both
from the perspective of public funding and collaborative
international projects?
See also: Agronomy of Grain Growing: Sorghum: Production and
Improvement Practices; Breeding of Grains: Barley: Genetics and
Breeding; Genetics of Grains: Rice: Genetics; Wheat Genetics and
Genomics; The Basics: Grain and Plant Morphology of Cereals and
how characters can be used to identify varieties; Taxonomic
Classification of Grain Species; The Cereal Grains: An Overview of
the Family of Cereal Grains Prominent in World Agriculture; Maize:
Overview; Millet Minor: Overview; Millet Pearl: Overview; Oats:
97
Overview; Rye Grain: Its Genetics, Production, and Utilization; Triticale:
Overview.
Further Reading
Clark LG and Pohl RW (1996) Agnes Chases First Book of Grasses. Washington, DC:
Smithsonian Press.
Elbaum R, Zaltzman L, Burgert I, and Fratzl P (2007) The role of wheat awns in the seed
dispersal unit. Science 316: 884886.
Doebley JF, Gaut BS, and Smith BD (2006) The molecular genetics of crop
domestication. Cell 127: 13091321.
Abbo S, Gopher A, Peleg Z, et al. (2006) The ripples of The Big (agricultural) Bang:
The spread of early wheat cultivation. Genome 49: 861863.
Allaby RG, Fuller DQ, and Brown TA (2008) The genetic expectations of a protracted
model for the origins of domesticated crops. PNAS 105: 1398213986.
Diamond J (2002) Evolution, consequences and future of plant and animal
domestication. Nature 418: 700707.
Tanno K-I and Willcox G (2006) How fast was wild wheat domesticated? Science
311: 1886.
Harlan JR (1992) Crops and Man, 2nd edn. Madison, WI: American Society of
Agronomy.
Zohary D, Hopf M, and Weiss H (2014) Domestication of Plants in the Old World: The
Origin and Spread of Cultivated Plants in South-West Asia, Europe and the
Mediterranean Basin, 4th edn. New York: Oxford University Press.
Badr A, Muller K, Schafer-Pregl R, et al. (2000) On the origin and domestication history
of barley (Hordeum vulgare). Molecular Biology and Evolution 17: 499510.
Morrell PL and Clegg MT (2007) Genetic evidence for a second domestication of barley
(Hordeum vulgare) east of the Fertile Crescent. PNAS 104: 32893294.
Van Heerwaarden J, Doebley J, Briggs WH, et al. (2011) Genetic signals of origin,
spread, and introgression in a large sample of maize landraces. PNAS
108: 10881092.
Mathur PN (2012) Global Strategy for the Ex Situ Conservation of Pearl Millet and Its
Wild Relatives. Rome, Italy: Global Crop Diversity Trust.
Harlan JR (1996) The tropical African cereals. In: Harris DR and Hillman GG (eds.)
Foraging and Farming: The Evolution of Plant Exploitation. London: Unwin
Hyman Ltd.
Loskutov IG (2008) On evolutionary pathways of Avena species. Genetic Resources and
Crop Evolution 55: 211220.
Yuan X, Xu Q, Dong G, et al. (2012) A map of rice genome variation reveals the origin of
cultivated rice. Nature 490: 497501.
Sackville-Hamilton R (2010) Global Strategy for the Ex Situ Conservation of Rice
Genetic Resources. Manila, Philippines: International Rice Research Institute.
Linares OF (2002) African rice (Oryza glaberrima): History and future potential. PNAS
99: 1636016365.
Shang H-Y, Wei Y-M, Wang X-R, and Zheng Y-L (2006) Genetic diversity and
phylogenetic relationships in the rye genus Secale L. (rye) based on Secale cereale
microsatellite markers. Genetics and Molecular Biology 29: 685691.
Mace ES, Tai S, Gilding EK, et al. (2013) Whole-genome sequencing reveals untapped
genetic potential in Africas indigenous cereal crop sorghum. Nature
Communications 4: 2320. http://dx.doi.org/10.1038/ncomms3320.
Marcussen T, Sandve SR, Heier L, et al. (2014) Ancient hybridizations among the
ancestral genomes of bread wheat. Science 345: 287-12287-15. http://dx.doi.org/
10.1126/science.1250092.
Kilian B, Ozkan H, Deusch O, et al. (2007) Independent wheat B and G genome origins
in outcrossing Aegilops progenitor haplotypes. Molecular Biology and Evolution
24: 217227.
Luven P (1988) Traditional Food Plants. FAO Food and Nutrition Paper 42. FAO.
Vietmeyer ND (1996) Lost Crops of Africa: Grains. vol. 1. Washington, DC: Board on
Science and Technology for International Development, National Research Council,
National Academy Press.
Wiersema JH and Leon B (2013) World Economic Plants: A Standard Reference, 2nd
edn. New York: CRC Press, Taylor & Francis Group.
Williams JT (ed.) (1995) Cereals and Pseudocereals London: Chapman and Hall.
Brkljacic J, Grotewold E, Scholl R, et al. (2011) Brachypodium as a model for the
grasses: Today and the future. Plant Physiology 157: 313.
Brutnell TP, Wang L, Swartwood K, et al. (2010) Setaria viridis: A model for C4
photosynthesis. Plant Cell 22: 25372544.
98 THE CEREAL GRAINS | Cereals: Domestication of the Cereal Grains
Relevant Websites
http://www.ars-grin.gov A searchable United States Department of Agriculture crop
germplasm database providing information on scientific and common names,
distribution, economic importance, and references.
http://www.ceg.icrisat.org/ipgsc.html International Pearl Millet Genome Sequencing
Consortium (global reference genotype Tift 23D2B1).
http://www.cgiar.org/cgiar-consortium/ Consultative Group on International
Agricultural Research (CGIAR) with five independent international centers conducting
basic and applied research on barley, maize, millet, rice, sorghum, and wheat.
http://www.croptrust.org/ A CGIAR partner with crop diversity programs targeting
conservation of the major cereal grains.
http://mansfeld.ipk-gatersleben.de An internet-searchable, version of Mansfelds
World Database of Agricultural and Horticultural Crops, which is set up for queries
on the taxonomy, common names, distribution, and other miscellaneous
information about crop plants.
http://www.public.iastate.edu/imagefpc/IBSC%20Webpage/IBSC%20Templatehome.
html International Barley Sequencing Consortium.
http://rgp.dna.affrc.go.jp/IRGSP/ International Rice Gene Sequencing Project (Oryza
sativa ssp. japonica cv. Nipponbare).
http://www.theplantlist.org/ Searchable database offering current listings of
accepted plant names and direct links to checklist data held by collaborating
institutions.
http://wheat.pw.usda.gov The United States Department of Agriculture searchable
database for a variety of topics dealing with the genomics, mapping, germplasm,
pathology, and taxonomy of barley, rye, triticale, wheat, and oats.
http://www.wheatgenome.org/ International Wheat Genome Sequencing Consortium
(Triticum aestivum).
 Maize: Overview
MP Scott and M Emery, Iowa State University, Ames, IA, USA
Published by Elsevier Ltd.
Topic Highlights
Maize is a member of the grass family Poaceae and the
closest known relative is the grassy weed teosinte.
Maize is a major crop, accounting for 32% of the total cereal
production worldwide in 2013.
Maize kernels provide nutrition in the form of starch, oil,
and protein.
Culinary dishes incorporating maize as an ingredient are
consumed worldwide.
Maize is processed via dry milling and wet milling and
nixtamalization.
Maize is a model organism in research.
Maize cultivars include inbred lines, single-cross hybrids,
double-cross hybrids, and open-pollinated varieties.
Biotechnology has allowed for reduced inputs and higher
outputs in maize production; however, some societies have
concerns regarding the use of genetically modified maize.
Learning Objective
To acquire a greater understanding of the role maize plays
in the world and how technology seeks to enhance maize
quality and profitability.
Introduction
Zea mays, more commonly referred to as maize, is a member of
the grass family Poaceae, or true grasses. Maize is thought to
have originated 5570 million years ago in what is now Cen-
tral or South America and has since diversified into nearly
10 000 nondomestic relatives. Figure 1 shows a phylogenetic
tree of grass species related to maize. There exists no direct
ancestor for maize; however, to date, the closest relative to
maize are the teosintes. Prehistoric selection has resulted in
ears lacking seed cases called glumes and seeds that adhere to
the cob until manual removal. These alterations limit the
ability of maize to survive without human intervention.
Maize is an annual plant with C4 metabolism, making it very
efficient at carbon fixation. It has the greatest global produc-
tion of any crop species around 800 million tonnes was pro-
duced worldwide in 2013, accounting for 32% of the total
cereal production. The top three producers include the United
States, China, and Brazil. Maize is grown on more areas of the
planet than any other crop and is grown on every continent
except Antarctica. The grain of maize is used for food, feed, and
industrial products including biodegradable foams, plastics,
and adhesives. Additionally, maize stover, the leaves and
stalk of the maize plant, is used for forage, biofuel production,
and chemical production.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00022-X
Maize Reproduction
Maize is a monoecious plant, meaning it has both male and
female reproductive organs on the same plant. Flowers mature
after
6070 days of vegetative plant growth. Male staminate
flowers develop into tassels and are found on the uppermost tip
of the main stem. Female pistillate flowers are found in one or
more ears located at nodes along of the stem. Typical maize
varieties are diploid, containing two sets of ten chromosomes.
Copious amounts of pollen (up to 1 billion grains per plant) are
shed from anthers and dispersed by air currents. While the
majority of the pollen falls close to the plant, a small portion
of the pollen can be carried over great distances via the wind.
Industry standards typically consider plants separated by a dis-
tance of 200 m to be reproductively isolated. Fertilization occurs
by the process of double fertilization common to angiosperm
species. A pollen grain carrying two nuclei lands on a silk and
germinates to produce a pollen tube. The pollen tube grows
down the length of the silk until it reaches the embryo sac
where it ruptures releasing the two sperm nuclei. The first
sperm cell fuses with the egg cell to produce the embryo, the
organ that ultimately develops into the next-generation plant.
The second sperm cell fuses with the central cell of the embryo
sac giving rise to the endosperm, the storage tissue that nour-
ishes the developing seedling until it is capable of living inde-
pendently. Grain fill to maturity takes about 40 days.
Maize Kernel Composition
The mature maize kernel is referred to as a caryopsis and is not
a true seed but rather a one-seeded fruit. Kernels are composed
of four organs: the pericarp, embryo, endosperm, and pedicel.
Physical properties, such as hardness, shape, size, color, and
composition, vary among maize varieties.
The main organs of a maize kernel are shown in Figure 2. The
outer layer of the kernel is the pericarp and encloses the kernel
for protection. The endosperm comprises the majority of the
kernels inner contents. The endosperm itself is composed of
four tissue types: the aleurone (outer) layer, the starchy endo-
sperm, the basal endosperm transfer layer, and the embryo-
surrounding region. The endosperm provides nutrients in the
form of sugars and amino acids to the growing embryo.
The embryo is composed of the following: the scutellum (the
monocotyledon that absorbs nutrients during germination), the
coleoptile (protective sheath of the emerging shoot), the plu-
mule (young plant), the radicle (primary root), and the coleo-
rhizae (protective sheath of emerging root). The tip cap serves to
attach the kernel to the cob and protect the kernel.
In terms of nutritive composition, the kernel can be further
classified into five main components. The typical number 2
yellow dent maize kernel contains
72% starch, 9.5% protein,
4.3% oil, 1.4% ash, and 2.6% sugar.
99
100 THE CEREAL GRAINS | Maize: Overview
Starch
Starch is the most abundant component in maize kernels and
serves as an efficient storage molecule for glucose. Starch accu-
mulates in the form of dense-insoluble granules. It is com-
posed of two main components: amylose and amylopectin.
Amylose is predominantly a linear polymer composed of 1,4-
linked a-D-glucan chains. In contrast, amylopectin is highly
branched by a-1,6-glycosidic bonds. Starch biosynthesis
requires the coordinated activities of a myriad of enzymes,
including starch synthases and starch branching enzymes and
starch debranching enzymes. Enzymatic activity within the
kernel alters starch content and properties. The degree of
branching and branch chain length are starch properties that
can vary considerably among maize varieties. Maturity also
affects starch quality. Traits sought in a commercial setting
include gel strength, viscosity, and thermal properties such as
gelatinization. Maize starch provides 4 cal g1.
Genetic mutations can confer altered starch phenotypes.
Mutant alleles of waxy1 (wxy1) produce 100% amylopectin
starch, which is useful as a thickening agent in foods. Mutation
of the amylose-extender (ae) gene leads to high-amylose starch
(HAS) with a range of amylose values from 25% to 80%. HAS
A phylogeny of diploid grass species
Barley
Goat Grass
Einkorn Wheat
Tall Fescue
Asian Rice
Teosinte
Maize
Eastern Gammagrass
Sorghum
Fountain Grass
Figure 1 A phylogeny of diploid grass species. Adapted from Gaut
BS, Le Thierry DEnnequin M, Peek AS, and Sawkins MC (2000) Maize as
a model for the evolution of plant nuclear genomes. PNAS 97:
70087015. http://dx.doi.org/10.1073/pnas.97.13.7008.
Figure 2 The mature maize kernel, showing component parts.
Reproduced from Encyclopedia of Grain Science.
is known for its slow digestion in vivo. The sugary-1 (su1) and
shrunken-2 (sh2) lead to kernel phenotypes that are sweeter
than field corn and are used to produce sweet corn varieties
for canning and fresh consumption.
Oil
Oil from a kernel of typical Corn Belt maize, number 2 yellow
dent, contains
62% linoleic acid, 25% oleic acid, 10%
palmitic acid, 2% stearic acid, and 1% linolenic acid; saturated
fatty acids equate to
12% of the total lipid content. The oil
within the maize kernel provides 9 cal g1. Linoleic, linolenic,
eicosapentaenoic, and docosahexaenoic fatty acids are shown
to have a positive correlation with cardiovascular health.
A ratio of 6:1, linoleic to linolenic, is recommended.
Similar to starch content and quality, studies demonstrate
that exotic germplasm possesses extensive ranges of fatty acid
composition. Exotic lines are crossed to yield varieties with
increased oil content. Oil content varies across inbred maize
lines and across varied environments. Total fatty acid compo-
sition varies throughout kernel development and ultimately
increases as the kernel matures. Oil content is believed to be
affected by a large number of loci and is a highly heritable trait.
Certain breeding schemes aim solely at increasing lipid content
and/or quality. Duvick (2003) altered fatty acid content by
introducing Tripsacum genes, a wild relative of maize, into
various maize lines.
Fatty acid stability is directly correlated to saturation level.
Linolenic is the least stable fatty acid, containing three points
of unsaturation. Oleic fatty acids are much more stable and less
prone to oxidation. Oleic fatty acids are monounsaturated.
Once oxidation begins, it cannot be stopped or reversed and
ultimately leads to rancidity.
Protein
Protein is another vital component in the maize kernel. Seed
proteins are divided into four classes: albumin, globulin, pro-
lamin, and glutelins. The major storage proteins in maize are
prolamins, also referred to as zeins. Eighty percent of the stored
protein in maize is found in the endosperm. Because of the
amino acid balance of zeins and their abundance in the endo-
sperm, lysine, tryptophan, and methionine are typically at low
levels in maize. Maize is therefore not a complete protein
source and must be eaten with complementary protein sources
to ensure requirements for the essential amino acids are met.
Many countries rely on maize as their main food source; in
turn, essential amino acid deficiencies such as kwashiorkor and
pellagra frequently occur. Maize protein provides 4 cal g1.
Research aims to increase the quality of protein in maize.
First observed in 1920, the Opaque-2 (o2) mutation causes a
decrease in the amount of zein content and thus a higher ratio
of nonzein proteins with increased levels of essential amino
acids. Unfortunately, this mutation results in reduced kernel
hardness, yield, and fungal and pest resistance. To overcome
this deficiency, modifier genes have been introduced into o2
varieties that increase kernel hardness. The resulting maize is
called quality protein maize and grown in many parts of the
world where it has contributed to improved nutrition. In addi-
tion to o2 mutants, Floury2 (fl2) mutants have shown to have
 THE CEREAL GRAINS | Maize: Overview 101

improved amino acid balance. Less abundant components of then used to coarsely grind the maize kernels. Several steps of
the maize kernel include fiber, minerals, vitamins, anthocya- size and weight separation, in addition to regrinding, yield
nins, and antinutrients. maize grits, flour, and fiber. The quality, content, and end use
of the maize must be considered before entering the wet- or
dry-milling process.
Maize kernels with a softer endosperm perform better in the
Maize in Food
wet-milling process. The wet-milling process entails steeping
maize in a dilute sulfur dioxide solution to soften the kernel
Maize is a food ingredient that brings commonality to culinary
and separate it into its smaller components. The germ can be
cultures across the world. Cultural traditions and corn varieties
first removed and later processed for oil. The remaining com-
dictate how maize is incorporated into a wide variety of foods.
ponents are ground and separated further into grits, flour, and
The main kernel components can be separated and processed
fiber. Further processing yields corn gluten meal, and starch.
into products such as corn starch for thickening and binding
Maize starch and high-fructose corn syrup are a main end
agents and corn oil for frying and baking; whole grain kernels
product of the wet-milling process in the United States.
are used in popped popcorn or ground into corn meal and
used in breads, biscuits, and cereals. From enchiladas, tamales,
totopos, tostaditas, and tortillas, virtually every Mexican dish
Maize Kernels
uses maize. Maize porridges are seen across the world: referred
to as puliszka and malderash in Hungary, posho in Africa,
polenta in Europe, grits in the United States, and kpekple in
Ghana. Maize meal can be ground and fermented into sora, a
Dry milling Wet milling Nixtamalization
maize beer in Peru, or used to make hard alcohols such as
whiskey and bourbon. Maize is truly a cross-cultural food.

Flaking frits Starch

Maize Processing Brewing grits Germ
Other grits Gluten Masa
Maize kernel quality and physical attributes determine its end Corn meal/flour Bran Dry masa flour
use. The United States recognizes five grades of maize and three Germ Steep water liquor
classes: yellow, white, and mixed maizes. Food maize typically Bran
specifies number 1 grade yellow or white dent corn (Figure 3).
Additionally, the manner in which maize is processed is a vital
component in its incorporation into foods (Figure 4).
Food industries
Harder kernels are desirable for storage, shipping, and han-
Breakfast cereals, snacks, baking, brewing, distilled spirits, oil,
dling; dry milling calls for a kernel with a harder endosperm sweeteners, corn and tortilla chips
void of cracks. Dry milling is often used to produce baked
goods, breakfast cereals, and ethanol. In the dry-milling pro- Figure 4 Maize food processing determines maize as food ingredient.
cess, tempering the grain is a vital first step. A hammer mill is Adapted from Encyclopedia of Grain Science.

U.S. Grades and Grade Requirements for Maize

Minimum Test, Maximum Percent Allowed
Weight/Bushel Damaged Kernels Broken Kernels and
Grade (Ib) Heat-Damaged Total Foreign Material
U.S. 1 56.0 0.1 3.0 2.0
U.S. 2 54.0 0.2 5.0 3.0
U.S. 3 52.0 0.5 7.0 4.0
U.S. 4 49.0 1.0 10.0 5.0
U.S. 5 46.0 3.0 15.0 7.0

U.S. Sample Grade:

(a) Does not meet the requirements for grades U.S. No. 1,2,3,4, or 5: or
(b) Contains stones which have an aggregate weight in excess of 0.1 percent of the sample
weight, 2 or more pieces of glass, 3 or more crotalaria seeds (Crotalaria spp.), 2 or more
caster beans (Ricinus communis L.), 4 or more particles of an unknown foreign
substance(s) or a commonly recognized harmful or toxic substance(s), 8 or more
cockleburs (Xanthium spp.) or similar seeds singly or in combination, or animal filth in
excess of 0.20 percent in 1,000 grams; or
(c) Has a musty, sour, or commercially objectionable foreign odor; or
(d) Is heating or otherwise of distinctly low quality.

Figure 3 The US maize grading scale. Reproduced from USDA (2013) Grain Inspection Handbook. Pamphlet. USDA Federal Gain Inspection Service,
Washington, DC.
102 THE CEREAL GRAINS | Maize: Overview
Nixtamalization, dating back from 1200 to 1500 BC, is an
ancient type of maize processing that includes rendering ker-
nels into a paste to increase the bioavailable nutrients such as
calcium and digestible iron. Kernels are steeped in a water/lime
solution over heat and ground into masa, also known as maize
dough, that is used to produce tortillas, corn chips, and other
food products.
Maize in Science
Maize is an important model organism in genetic research. It
has several attributes that make it attractive for this purpose. It
is a large plant and phenotypic analyses are easily done. Each
plant produces an ear typically containing 100400 kernels. It
is broadly adapted and has tremendous genetic diversity. Maize
has a moderately sized genome of
2.5 gigabase pairs. A vast
collection of mutant stocks have also been developed that
assist in research; this has allowed for many genes to first be
characterized molecularly in maize. Being a diploid species,
genetic manipulation and analysis is less complex than in
species with a higher ploidy level. Additionally, the large phys-
ical size of the maize chromosomes is a great benefit to cyto-
genetic researchers.
Research on maize has led to several key discoveries. Per-
haps most notable is the discovery of transposons by Barbara
McClintock, for which she was awarded the Nobel Prize in
Physiology or Medicine in 1983. Cytogenetic studies in maize
resulted in an understanding of genetic recombination and
enabled genetic mapping. The role of telomeres was deter-
mined in maize. Through collaborative efforts, it was one of
the first crops to have its genome completely sequenced.
Maize Breeding, Genetics, and Biotechnology
Maize Cultivar Types
A cultivar is a plant variety that has been developed for a
specific use. Several types of maize cultivars are grown includ-
ing inbred lines, single-cross hybrids, double-cross hybrids,
and open-pollinated varieties (Figure 5). Inbred lines are cre-
ated by successive generations of self-pollination. The resulting
plants are genetically homozygous and phenotypically
Single cross Inbred line  Inbred line
A B
Modified single Inbred line  Inbred line  Inbred line
cross A1 A2 B
Three-way cross Inbred line  Inbred line  Inbred line
A B C
Double cross
Inbred line  Inbred line  Inbred line  Inbred line
A B C D
Figure 5 Types of hybrids grown commercially in North America.
Reproduced from Encyclopedia of Grain Science.
homogeneous. Due to inbreeding depression, inbred lines
have low yield and are not used for grain production. Their
main purpose is in the production of hybrid seed. When two
inbreds are cross-pollinated, a single-cross hybrid results.
Single-cross hybrids are genetically heterozygous and pheno-
typically uniform. Because of the difficulties of producing seed
on inbred lines, several types of hybrids have been developed.
An open-pollinated variety is a population of plants that is
genetically heterozygous and phenotypically nonuniform. As
the name implies, the seed of open-pollinated varieties is pro-
duced by allowing natural pollination to occur in the popula-
tion. Synthetic populations are derived from intermating
several varieties that are frequently used in breeding programs
to produce inbred lines.
Mechanized agriculture has led to a preference for hybrids
because of their uniformity and high yields. The process of
hybrid improvement and seed production has become highly
industrialized. Industrial maize breeding has led to greatly
increased yields. Open-pollinated varieties require much less
infrastructure for seed production and genetic improvement
and are often grown in developing countries.
Hybrid Maize Breeding
Hybrid maize breeding allows breeders to capture and fix
extremely productive genotypes by taking advantage of hybrid
vigor. Productivity and vigor in maize plants are generally
proportional to the degree of heterozygosity. Thus, inbred
lines, although uniform and reproducible, are usually poor
agronomic purposes. Heterozygosity and performance can be
restored by crossing unrelated inbred lines to make hybrids.
Inbred lines are classified into heterotic groups according to
their ability to form productive hybrids in combination with
other groups and their suitability as a male or female parent.
For example, nearly all inbreds used as females in North
American hybrids are in the Stiff Stalk heterotic group. Devel-
opment and maintenance of inbred lines and testing hybrid
combinations require a great deal of infrastructure and exper-
tise that are not available to most farmers or even small seed
companies and are therefore largely done by large seed
companies.
Uniformity is essential in efficient and profitable produc-
tion of maize. Superior technology and machinery has assisted
with such uniformity. Improved accuracy in the evaluation of
cultivars has allowed for large genetic gains and the overall
creation and advancement of superior maize inbreds. Superior
farm equipment equipped with GPS and computer monitoring
systems has led to optimal planting depth, density, and spacing
and precise measurements of grain yield during harvest. In the
future, precision agriculture will continue to increase produc-
tivity by optimizing inputs, such as maize variety and fertilizer
amount, on a per land area basis.
Maize Biotechnology
Biotechnology gives the ability to introduce genes from any
source into the maize genome. Two types of traits derived from
genetic engineering methods are currently in commercial pro-
duction: insect resistance and herbicide tolerance.

Insect-resistant maize decreases the need of pesticide appli-
cations directly to the plant. The use of pesticides in the United
States has been reduced to 6% since 1996, a total of 172
million kilograms per year. Fewer tonnes of chemical are
applied, benefiting the health of the environment and proving
economically beneficial for the farmer. From 1996 to 2010, the
income of US farmers increased to a total of $21.7 billion
dollars, 23% of that profit was derived from 2010 alone. The
percentage of genetically engineered (GE) maize has increased
almost fourfold in 12 years (Figure 6.). The US Department of
Agriculture (USDA) Economic Research Service reports that
Bacillus thuringiensis (Bt)-resistant maize decreased the amount
of insecticides per planted acres of Bt maize by 8% in the
United States. Herbicide-tolerant maize is amenable to low to
no-till agriculture. Minimal to no tillage results in decreased
fuel usage and reduction of greenhouse gas emissions, as well
as less soil compaction and erosion. Additionally, crop residue
left on top of the soil increases the levels of organic carbon
sequestration. Soil and water quality are increased due to
decreasing soil erosion and nutrient loss (Committee on the
Impact of Biotechnology on Farm-Level Economic and
Sustainability and National Research Council, 2010; National
Research Council, 2010).
Of the 159 million hectares of maize grown globally in
2012, 55.1 million hectares (35%) was genetically modified
(GM) maize. Legislation regulating such crops varies among
countries. The United States regulates genetically modified
organisms based on the end product. Three groups with differ-
ing perspectives and expertise regulate GM crops in the United
States: the US Environmental Protection Agency, the Food and
Drug Administration (FDA), and USDA. GM crops must be
verified free from environmental and human toxins and for-
eign proteins deemed allergenic. The FDA policy established in
1992 considers the currently approved GM crops to be sub-
stantially equivalent to non-GM crops and deemed generally
GE Maize in the Unites States
100
90
80
Percent GE Maize
70
60
50
40
30
20
2000 2002 2004 2006 2008 2010 2012
Year
Figure 6 Percentage of all maize grown in the United States that is
genetically engineered (GE). Reproduced from USDA (2014) Adoption of
genetically engineered crops in the U.S.
THE CEREAL GRAINS | Maize: Overview 103
recognized as safe under the Federal Food, Drug, and Cos-
metic Act; therefore, foods made with approved GM varieties
do not require premarket approval. Acceptance of GM maize
by the consumer varies by country. The European Union (EU)
regulates GM crops based on the process in which they are
produced. The EU tends to be cautious of GM crop consump-
tion. The British Press often refers to such crops as Franken-
stein Foods. Protester of third-world countries have been
known to destroy entire fields of GM crop despite starvation
in the country. The major concerns over the production of GM
maize are pollination of weedy species or non-GM maize by
GM pollen resulting in undesirable transfer of the transgene
and the impact of transgenes on nontarget species, particularly
beneficial insects, and the development of insects resistant to
the mode of action of the insecticidal transgenes in use.
Researchers and regulatory agencies continue to develop new
deployment strategies in an effort to minimize these risks.
Exercises for Revision
How has human selection led to the evolution of maize?
What countries are the top three producers of maize in the
world?
What kernel characteristics must be taken into consider-
ation when processing? Why?
What characteristics make maize a good model organism?
Imagine a scenario where a GM maize line is created that
corrects a serious nutritional deficiency; however, the pol-
len of the maize line is known to be compatible with weeds
grown in the area where the nutritional deficiency is found.
Should this maize line be released in this area? Explain your
reasoning.
Does GM maize support or hinder progression toward a
sustainable agriculture?
Exercises for Readers to Explore the Topic Further
Research continually strives to produce maize that requires
fewer inputs and produces higher outputs while maintaining
kernel quality, yield, and nutritive composition. The creation
of new maize lines with novel traits will continue to be seen.
What novel traits do you think should be introduced?
Continued improvements in breeding and biotechnology
will allow the development of maize varieties with
improved yield and agronomic traits. What do you think
are the practical limits to yield improvement.
Educating consumers about the food they consume and
advocating individuals to take an active interest in the
production of their food will be helpful in ensuring sus-
tainable food production. Design educational material for
maize-based food in your community.
See also: Barley, Rice and Maize Processing: Maize: Dry
Milling; Maize: Wet Milling; Oil from Rice and Maize; Carbohydrates:
Resistant Starch and Health; Starch: Starch Architecture and Structure;
Starch: Synthesis; Food Grains and the Consumer: Genetically
Modified Grains and the Consumer; Genetics of Grains: Maize:
104 THE CEREAL GRAINS | Maize: Overview
Genetics; Maize: Other Maize Mutants; Non-wheat Foods: Maize:
Foods from Maize.
Further Reading
Brookes G and Barfoot P (2005) GM crops: The global economic and environmental
impact-the first nine years 19962004. AgBioforum 8: 187196.
Campbell MR, White PJ, and Pollak LM (1994) Dosage effect at the sugary-2 locus on
maize starch structure and function. Cereal Chemistry 71: 464468.
Corn Breeding: Types of Cultivars (2014) from http://passel.unl.edu/pages/
informationmodule.php?idinformationmodule1099683867&topicorder8&
maxto9&minto1.
Crawley MJ, Brown SL, Hails RS, Kohn DD, and Rees M (2001) Transgenic crops in
natural habitats. Nature 409: 682683.
Dudley JW and Lambert RJ (1992) Ninety generations of selection for oil and protein in
maize. Maydica 37: 17.
Duvick D (1996) Plant breeding, an evolutionary concept. Crop Science 36: 539548.
Eckhoff SR (2004) Wet milling. In: Wrigley C (ed.) Encyclopedia of Food Grains, 1st ed.,
pp. 225241. Oxford: Academic Press.
Flint-Garcia SA, Bodnar AL, and Scott MP (2009) Wide variability in kernel
composition, seed characteristics, and zein profiles among diverse maize inbreds,
landraces, and teosinte. Theoretical and Applied Genetics 119: 11291142.
Gaut BS, Le Thierry DEnnequin M, Peek AS, and Sawkins MC (2000) Maize as a model
for the evolution of plant nuclear genomes. PNAS 97: 70087015. http://dx.doi.org/
10.1073/pnas.97.13.7008.
Hallauer AR (2004) Specialty corns. In: Corn: Origin, History, Technology, and
Production, pp. 897933. Hoboken, NJ: Wiley.
Hannah LC (2005) Starch synthesis in the maize endosperm. Maydica 50: 497506.
Jellum MD (1970) Plant introductions of maize as a sourced of oil with unusual fatty
acid composition. Journal of Agricultural and Food Chemistry 18: 365370.
Jennings S, Myers D, Johnson L, and Pollak LM (2002) Effects of maturity on corn
starch properties. Cereal Chemistry 79: 703706.
Jeon J, Ryoo N, Hahn T, Walia H, and Nakamura Y (2010) Starch biosynthesis in cereal
endosperm. Plant Physiology and Biochemistry 48: 383392.
Kiesselbach TA (1949) The structure and reproduction of corn. Lincoln, NE: University
of Nebraska.
Lorenz A, Scott P, and Lamkey K (2008) Genetic variation and breeding potential of
phytate and inorganic phosphorus in a maize population. Crop Science Society of
America 48: 7984.
McClintock B (1950) The origin and behavior of mutable loci in maize. Genetics
36: 344355.
Mertz ET, Bates LS, and Nelson OE (1964) Mutant gene that changes protein
composition and increases lysine content of maize endosperm. Science
145: 279280.
Nelson OE, Mertz ET, and Bates LS (1965) Second mutant gene affecting the amino acid
pattern of maize endosperm proteins. Science 150: 14691470.
Orthoefer FT and Eastman J (2004) Corn processing and products. In: Corn: Origin,
History, Technology, and Production, pp. 867896. Hoboken, NJ: Wiley.
Pollak LM and Scott MP (2005) Breeding for grain quality traits. Maydica 50: 247257.
Poneleit CG and Davis DL (1972) Fatty acid composition of oil during maize kernel
development. Phytochemistry 11: 34213426.
Prasanna BM, Vasal SK, Kassahun B, and Singh NN (2001) Quality protein maize.
Current Science 81: 13081319.
Rooney LW, McDonough CM, and Waniska RD (2004) The corn kernel. In: Smith CW,
Betran J, and Runge ECA (eds.) Corn: Origin, History, Technology, and Production,
pp. 273303. Hoboken, NJ: Wiley.
Scanlon MJ and Takacs EM (2009) Kernel biology. In: Bennetzen JL and Hake SC (eds.)
Handbook of Maize. New York, NY: Springer ScienceBusiness Media.
Sema-Saldivar SO (2004) Foods from maize. In: Wrigley C (ed.) Encyclopedia of Food
Grains, 1st ed., pp. 242253. Oxford: Academic Press.
Snow AA (2002) Transgenic crops: Why gene flow matters. Nature Biotechnology
20: 542.
Strable J and Scanion M (2013) Maize (Zea mays): A model organism for basic and
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Tucker J (2011) U.S. Regulation of Genetically Modified Crops. Case Studies in
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dualuse-agriculture/2-agricultural-biotechnology/us-regulation-of-genetically-
engineered-crops.html
USDA (2013) Grain Inspection Handbook. Pamphlet. USDA Federal Gain Inspection
Service, Washington, DC.
USDA (2014) Adoption of genetically engineered crops in the U.S., 2014.
Vasal SK (2000) High quality protein corn. In: Hallauer AR (ed.) Specialty Corns, 2nd
edn. Boca Raton, FL: CRC Press.
Velu V, Nagender A, Prabhakara Rao PG, and Rao DG (2006) Dry milling characteristics
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Vineyard ML and Bear RP (1952) Amylose content. Maize Genetics Cooperation
Newsletter 26: 5.
Vogel K and Burson B (2004) Breeding and Genetics. Madison, WI: American Society of
Agronomy, Crop Science Society of America, Soil Science Society of America,
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Wijendran V and Hayes KC (2004) Dietary n-6 and n-3 fatty acid balance and
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10.1146/annurev.nutr.24.012003.132106.
Wilkes G (2004) Corn, strange and marvelous: But is a definitive origin known?
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genetic model generates high-resolution mapping of QTL for protein quality in
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States of America 99: 1128111286.
 Wheat: An Overview of the Grain That Provides Our Daily Bread
CW Wrigley, University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Wheat is unique in its suitability for bread production
because of its gluten protein.
Bread has a major place in world language and culture
because of its importance in our diet.
Wheat, a member of the cereal family, is a monocotyledon-
ous grass of the genus Triticum. There are two species
common (bread) wheat and durum (suited to pasta).
Wheat is grown in a wide range of environments, including
tropical and very cold regions.
Worldwide wheat production is equivalent to about 300 g
of wheat per day for each person on Earth.
Wheat-based breads include leavened loaves, Arabic and
Middle Eastern flatbreads, and Chinese steamed breads.
Wheat has many other food uses, including noodles, pasta,
pastry, donuts, cakes, and cookies.
Nonfood uses include animal and fish feeds, starchgluten
manufacture, sweeteners, adhesives, and biofuels.
The main component of wheat flour is starch, followed by
protein, which is largely the gluten-forming gliadins and
glutenins. The fat content is low. Wheat is a major source of
energy, fiber, vitamins, and minerals in our diet.
Some individuals are intolerant to wheat in the diet. The
best characterized intolerance is celiac disease.
Learning Objective
To achieve understanding of wheat as a grain species, includ-
ing its origins, genetics, production, utilization, and its value in
our diet.
Introduction
Wheat is unique among the food grains. Only the wheat grain
contains storage proteins that form into a cohesive dough after
milling, adding water, and mixing. Only the dough from wheat
flour has the extensibility and elasticity to allow the growth of gas
bubbles (generally air and carbon dioxide) and to allow the
retention of the bubble structure throughout the bread baking
process. Wheat proteins dough-forming ability also accounts for
its value for making the wider range of foods, including noodles,
pasta, pastry and pizza crust, cakes, cookies, and even candies.
These are the reasons for the success of wheat not only as one of
the three most popular grains, together with rice and maize, but
also as the grain with the most stringent quality specifications.
Wheat, Bread, and the Human Race
Despite the wide range of uses of wheat, it is the word bread
that epitomizes the use of wheat. Bread and wheat are central
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00020-6
in the symbol of the FAO (Food and Agriculture Organization
of the United Nations); a wheat ear is the logo and the motto in
Latin Fiat panis, which means Let there be bread (Figure 1).
Likewise, the word bread has entered our everyday languages
as a symbol of food more generally.
Bread is international, but in its variations, there are
national characteristics, as illustrated in Figure 2. In the lives
of Polish and Russian people, bread (chleb) is a sacred item.
In such cultures, an honored guest is welcomed by the presen-
tation of a loaf of bread, covered with a lace cloth on a platter.
In Iran, sweet bread (Persian naan) may be taken as a gift
when visiting friends. In the Jewish Passover celebration, the
bread must be unleavened, as a reminder of the Israelites
hurried departure from slavery in Egypt, when there was no
time to leave the bread to rise overnight. From Jewish and
Christian cultures, various bread-related expressions have
entered everyday English: Cast your bread upon the waters,
Give us this day our daily bread, and Man shall not live by
bread alone. Similar expressions are found in other languages,
cultures, and religions.
Wheat is even more than bread, because its dough-forming
properties suit its use for the further range of foods in Figure 3
crackers, cookies, pastries, and noodles of many types
(Figure 4). Pasta types (such as spaghetti, macaroni, and taglia-
telle), distinct from noodles, are correctly made from durum
wheat (Figure 5). In addition, wheat is an important feed source
for animals, birds, and fish. Industrial uses include the separa-
tion of starch from gluten protein. The starch fraction is used for
paper, adhesives, and even to produce ethanol as a fuel. Gluten
is used to augment dough quality in baked goods and as an
additive to a wide range of foods.
The Greek goddess Demeter (depicted in Figure 6 with
awned wheat heads and stalks) was probably adopted and
renamed Ceres by the Romans, thus providing the origin of
our word cereal, which now includes all the grasslike grains,
for example, barley, rye, rice, and maize.
Wheat Species
The ancestors of modern wheats still grow in the regions where
they had their origins in the Fertile Crescent (Egypt through
Syria to the TigrisEuphrates valley) and further eastward. They
include the diploid einkorn wheats (Triticum boeoticum and
T. monococcum) and the tetraploid emmer wheats (T. dicoccoides
and T. dicoccum). Hexaploid bread wheat arose from natural
hybridizations between these tetraploid wheats and the dip-
loid species T. tauschii.
There are two modern wheat species: bread (or common)
wheat, T. aestivum, and durum wheat, T. durum (also desig-
nated T. turgidum ssp. durum), which is less common and is
mainly used for pasta and whole-grain uses such as couscous.
Figure 7 shows stands of common wheat almost ready to
105
106 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
Figure 1 The logo of the Food and Agriculture Organization uses bread
(panis) as a symbol of food in general. Reproduced with FAO
permission.
Figure 2 Bread types from various nations and cultures. Reproduced
with the permission of Perten Instruments.
Figure 3 The wider range of foods made from wheat flour. Reproduced
with the permission of Perten Instruments.
Figure 4 The many types of wheat-based noodles. Reproduced with the
permission of AWB Ltd.
Figure 5 Pasta types, such as spaghetti, are made from durum wheat, a
distinct species from bread wheat.
harvest. The most obvious distinction between these crops is
that one is an awned (bearded) variety and the other is not.
Not evident in the photos, however, are important distinctions
that indicate the suitability of the mature grain for the various
uses of wheat grain, such as protein content, grain hardness,
and potential for dough quality needed for specific food types.
Common wheat has 21 pairs of chromosomes, arranged in
three genomes (A, B, and D), reflecting the three original
diploid progenitors (seven pairs each) of hexaploid wheat.
The chromosomes are numbered 1A to 7A, 1B to 7B, and 1D
to 7D. Durum wheat, in contrast, has only 14 pairs of chro-
mosomes, numbered 1A to 7A and 1B to 7B, reflecting in turn
its origins. The genomes, and the chromosomes of correspond-
ing number (e.g., 1A vs. 1B vs. 1D), are said to be homoeolo-
gous that is, they are similar but not homologous (identical).
So, for example, chromosomes 3A, 3B, and 3D have homo-
eologous genes for red grain color.
 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
Figure 6 The Greek god of wheat, Demeter, drawn originally from a
painting found in Pompeii. Reproduced from Buller AHR (1919) Essays
on Wheat, p. 281. New York: McMillan Company.
Wheat Production and Utilization Worldwide
On average, world production of wheat totals about 700 mil-
lion tonnes (Mt) annually, grown on some 220 million hectares
(Mha), with a world average yield of 3.2 tonnes ha 1. Table 1
lists the countries with an annual production of over 3 million
tonnes. The European Union appears at the top as the largest
wheat producer; the contributing EU countries are listed indi-
vidually. The total cereal grain production is also listed for some
countries, indicating the relative importance of wheat com-
pared with other grains, especially rice and maize. Countries
such as Indonesia, Thailand, Mexico, and South Africa are
significant grain producers, but wheat is not their main crop.
The single-season data in Table 1 fail to show the fluctua-
tions in yield that occur from season to season. These year-to-
year changes are evident in the 5-year data in Table 2, but even
in this case, fluctuations are less obvious due to the averaging
effects of summing the production of many countries.
The ideal wheat-growing climates are subtropical, with
mean summer temperatures of about 15  C and an annual
rainfall of 300700 mm falling more in spring (during grain
filling) than in summer (providing opportunity for a dry har-
vest). Nevertheless, the climates in which wheat can be grown
vary greatly, as is exemplified by wheat growing in the far north
of America versus Mexico.
In northern parts of Europe and the United States, winter
wheats are sown in autumn, to lie under a snow cover through-
out winter, ready for the warmth of spring to bring them back
into growth and grain production. This vernalization process is
107
essential to the ongoing development of the plants into head-
ing, pollination, and grain development. Distinct varieties
(winter genotypes) of wheat are needed to suit these climatic
conditions.
On the Canadian prairies and the northern US plains
(North Dakota and Montana), the winters are too harsh to
permit the growing of the higher yielding winter-habit wheats.
In these very cold regions, spring wheats are sown after the
snow has melted, as soon as it is possible to access the fields in
spring. Thereafter, there is very rapid growth, permitting the
grain to mature for harvest just before the snows of autumn.
Harvesting can thus be difficult sometimes as the grain does
not finish its maturity and drying out in the head. In this case,
the crop is swathed cut down and left to dry prior to harvest-
ing (Figure 7(c)). By contrast, wheat growing in Kansas, the
United States, and down into Mexico may involve heat stress in
the late stages of grain filling, coming into summer. Wheat
growing in parts of Mexico and the Indian subcontinent
involves regions of high altitude.
Average wheat yields vary considerably between the coun-
tries in Table 1. Some of the highest yields are in European
countries where their intensive agricultural practices combine
with adequate rainfall. In some regions, where high inputs are
possible, there are ten-ton clubs, involving groups of growers
who regularly achieve yields of over 10 tonnes ha 1. By con-
trast, regions of dryland farming may reckon on little more
than 1 tonnes ha 1 (Table 1). Drought and disease may reduce
yields in either of these extreme cases.
Todays world average yield of about 3 tonnes ha 1
(Table 2) contrasts with historical yields, which was only
1 tonnes ha 1 at the end of the 1950s (Figure 8). This three-
fold increase has been partly due to improved agricultural
practices and especially because of the introduction of wheat
varieties with improved yield traits. Figure 8 also shows that
there have been only minor increases in recent decades in the
area sown to wheat worldwide.
In temperate regions, dual-purpose wheats may be sown.
These are usually late-maturing varieties sown early so that
stock may graze the early leaves but allow the grazed plants
to recover and produce grain.
The times of year for sowing and harvesting vary around the
world (Table 3). Harvest generally coincides with summer,
thus reflecting whether the particular country is in the northern
or southern hemisphere. Other factors include sowing date and
the maturity of the particular variety sown, plus constraints
such as avoidance of extremes of climate both very cold and
very hot.
The choices of sowing date and the maturity of the variety
sown are often challenging decisions. Rough calculations may
be made about the critical time of flowering. Very early flower-
ing brings the risk that a late frost will prevent proper flowering
with the consequence that many florets may be sterile. Late
flowering risks heat stress during grain filling with the conse-
quences of reduced grain yield and weakening of the dough
quality of the mature grain.
Much of wheat production is used in the countries in which
it is grown, but world trade accounts for about 20% of total
production, a higher proportion than for cereal grains more
broadly (Table 2). World wheat production is equivalent to
about 300 g of wheat per day for each person on Earth. This
108 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
Figure 7 Wheat in the field: (a) unawned wheat, (b) awned wheat, and (c) wheat that has been swathed to hasten the drying of the grain ready for
harvesting.
statistic suggests that there should be no problem with hunger.
However, about one-third of world wheat production is used
for a combination of seed use, animal feed (about 20%), and
industrial processing (about 10%). Furthermore, the places of
excess production are far from the places of human need.
Many countries produce more wheat than their domestic
requirements, resulting in world trade of about 140 million
tonnes annually. The main wheat-exporting nations are the
United States, Canada, Europe, Australia, and Argentina. Recip-
ient countries that import grain include Brazil, Egypt, Japan,
South Korea, Indonesia, Iran, and Iraq. Such international trade
and transport over thousands of kilometers contrast with the
subsistence farming of wheat in some regions, where farm-
grown grain is milled locally for the farm family.
The Grain That Becomes Our Food
The wheat grain is the result of internal fertilization of the
wheat flower. The glumes that enclosed the flower can be
seen in Figure 9 in which other florets have been removed
from the rachis (the backbone of the wheat head) to show how
the mature grain is held within the protecting outer glumes,
lemma, and palea (the husks of the mature wheat head). At the
time of fertilization, anthers released pollen to fertilize the
ovum, resulting in a grain, which grows through the series of
anatomical stages shown in Figure 10. The diagram in
Figure 10(c) corresponds to the transmission electron micro-
graph in Figure 11.
The lowest third of Figure 11 is the outer part of the endo-
sperm, showing cell walls that enclose starch granules (both
large (A-type) and small (B-type)). These outer endosperm
cells have a larger proportion of storage protein (forming the
matrix between the starch granules) than the inner endosperm.
Enclosing the endosperm is a layer of aleurone cells; beyond
that are the bran layers, which are shriveled in the mature
grain, having served their functions for the developing grain
(as seen in Figure 10).
It is the aim of the flour miller to remove these outer layers
and the germ, to produce white flour made up of the endo-
sperm, and then to reduce the endosperm to a fine powder.
Moisture is used to render the bran layers leathery so that they
leave the grain during milling as large flakes that are readily
sieved from the endosperm particles. Flour milling involves
repeated passes between rolls, first fluted and later smooth,
with sieving steps between the rolling. From one stage to
another, the flour stocks are conveyed pneumatically through
the complex system of pipes shown in Figure 12.
Grain hardness is a significant factor to the miller. The
endosperm of a soft wheat has air pockets that render the intact
 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread 109

Table 1 Wheat production by countries

Country/region Wheat production (Mt) Wheat yields (tonnes ha 1) Production of all cereal grains (Mt)

The European Union 143 5.5 304

China 121 5.0 480
India 93 3.1 240
The United States 58 3.2 436
Russia 52 2.2 88
France 39 7.2
Canada 33 3.2 59
Australia 26 1.9 37
Germany 25 8.0
Pakistan 24 2.8 34
Ukraine 22 3.4 60
Turkey 18 2.3 32
Kazakhstan 16 1.2
Iran 15 2.1
The United Kingdom 12 7.5
Argentina 11 3.0 47
Egypt 9 6.3
Poland 9 4.4
Spain 8 3.6
Italy 7 3.9
Morocco 7 2.1
Romania 7 3.6
Uzbekistan 7 4.9
Bulgaria 5 4.1
Denmark 5 6.8
Hungary 5 4.7
Afghanistan 4 1.6
Syria 4 2.7
World 706 3.2 2432

Data from the United States Department of Agriculture.

Table 2 Production and trade involving wheat (and all cereals) over five recent years

Wheat 2009/10 2010/11 2011/12 2012/13 2013/14a

Production 686 654 702 660 711

Utilization 656 658 699 684 695
Trade 131 120 147 140 142
Yield (tonnes ha 1) 3.15 3.03 3.21

All cereals 2009/10 2010/11 2011/12 2012/13 2013/14a

Production 2266 2257 2353 2305 2500

Utilization 2234 2271 2326 2323 2413
Trade 278 289 319 309 318
a
Estimate.

endosperm opaque and whitish. On crushing, the endosperm
of a soft wheat readily disintegrates into a fine powder, with
little damage to the starch granules.
In contrast, the intact endosperm of a hard wheat or durum
grain is translucent and vitreous. The starch granules are held
firmly in the matrix of storage protein. More energy must be
put into milling a hard grain, with the initial mill pass produc-
ing a coarse powder (semolina for durum), which must be
further reduced with repeated milling.
Starch damage results; some granules may be split into pieces.
In any case, some of the protein matrix will remain on the surface
of the granules, as is shown by the rough surface of the large A
granule in Figure 13. A small B granule was caught on top of the A
granule in the process of sample preparation. The scanning elec-
tron micrograph thus shows the contrast in size of the two types of
granule. A granule from a soft flour would have a smooth surface.
Starch damage is desirable for most bread making as it
facilitates access by starch-degrading enzymes (amylases) to
110 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread

700

600 Area (Mha)

Production (Mt)
500

400

300

200

100

0
61 964 967 970 973 976 979 982 985 988 991 994 997 000
19 1 1 1 1 1 1 1 1 1 1 1 1 2
Figure 8 Worldwide wheat production compared to area sown. Reproduced with permission from Marshall DR, Langridge P, and Appels R (2001)
Wheat breeding in the new century Preface. Australian Journal of Agricultural Research 52: iiv.

Table 3 Wheat harvesting dates

Country Hemisphere Harvest dates

Argentina South NovemberJanuary

Australia South OctoberJanuary
Canada North JulySeptember
China North MaySeptember
England North AugustSeptember
France North JuneJuly
India North FebruaryMay
Italy North JuneJuly
Russia North JulySeptember
The United States North MaySeptember

the starch substrate and increases water absorption. The lower

absorption of water for soft wheats is desirable for cookie and
cake manufacture on the other hand. Figure 14 illustrates the
result of amylase action on a starch granule, which may have
had its surface damaged during milling to provide ready access
Figure 9 A spikelet of wheat with other spikelets removed from the
by the enzyme. The layered crystalline structure inside the rachis (backbone) of the head. The lowest floret (at right) has been
granule can be seen. opened to reveal a grain lying between the glumes (husks lemma
and palea). The lemma bears a short awn. There are further florets to
the left and above.

Dough Formation and Baking

The formation of suitable gluten structure in dough requires
both the addition of water and mixing action. The course of
dough development can be followed by equipment used to
analyze dough quality, such as the mixograph (Figure 15). The
broad trace of mixing action reflects the movement of the bowl
containing the dough. From little initial resistance to mixing,
the progressive dough formation is shown by the rise of the
curve until it reaches a maximum. Beyond the peak, resistance
becomes less. Usually, a dough mixed to peak development is
most desirable for bread baking.
During the mixing process, the grains storage protein
hydrates and coalesces to form gluten strands that bind
together forming an elastic matrix holding the starch granules
and the gas cells. The strands of gluten are clearly seen in the
scanning electron micrograph of mixed dough in Figure 16.
Also visible are the gaps of gas cells and the larger (A-type)
starch granules. The smaller B-type granules are covered by the
gluten strands.
Bread making can involve a simple combination of flour,
water, salt, and yeast, with the mixed dough left for yeast action
to produce bubbles of carbon dioxide prior to baking. A series
 STYLE
PERICARP

PERICARP
OUTER INTEGUMENT
INNER INTEGUMENT
ANTIPODAL OUTER INTEGUMENT
EMBRYO SAC

THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
ENDOSPERM INNER INTEGUMENT
NUCELLUS VASCULAR BUNDLE
CREASE
EMBRYO NUCELLAR EPIDERMIS

FUNICULUS

(a) 0.8 mm
EMBRYO SAC
0.5 mm
(b) DEGENERATING NUCELLUS

ALEURONE
CELLS

PERIPHERAL
PERICARP
ENDOSPERM
ALEURONE
CELLS PRISMATIC
ENDOSPERM
PERIPHERAL
ENDOSPERM
CENTRAL
PRISMATIC ENDOSPERM
ENDOSPERM
VASCULAR
BUNDLE

(c) CREASE

111
Figure 10 Diagrams of the wheat grain through stages of development. (a) The wheat ovule prior to fertilization. (b) The developing grain. (c) The mature grain.
112 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
Figure 11 Transmission electron micrograph of the almost mature
grain, showing endosperm cells with starch granules.
Figure 12 The many pipes of the pilot flour mill at Kansas State
University carry the various flour stocks pneumatically from one stage of
the milling process to another.
of loaves resulting from this simplest procedure (Figure 17)
shows that different flour samples can produce a variety of loaf
sizes from the same starting amount of dough, the loaf with the
largest volume (height) being considered the best.
In practice, the baking of leavened breads (Figure 2) is
more complex, as is described in relevant articles in this ency-
clopedia. In addition, bread comes in many different forms.
There is a wide family of flatbreads made in the Middle East
and through the Indian subcontinent. Indian chapati bread is
illustrated in Figure 18. The baking of both chapatis and Arabic
Figure 13 Scanning electron micrograph of wheat starch granules
(A-type [large granule] and B-type).
Figure 14 Scanning electron micrograph of a wheat starch granule that
has been eroded by amylase enzymic action, revealing the crystalline
layers of internal structure.
flatbreads (classed as hearth breads, Figure 19) takes place
rapidly on a very hot surface such as a flat hearth or the side
of an oven. It is desirable for the two crusts of an Arabic bread
to separate during baking so that fillings may be inserted
before eating. These breads thus come from the oven looking
like an inflated balloon (Figure 19).
Yeast-leavened breads may be heated by means other than
an oven. Chinese steamed breads have steam as the means of
heating. For bagels, the initial form of heat is boiling water; a
ring of yeasted dough is briefly boiled and then baked, pro-
ducing a chewy texture. On the other hand, boiling oil is the
means of heating for a doughnut (Figure 20), which may be
 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread 113

400
Breakdown

Bandwidth
Resistance

at peak

200

Peak
resistance
Mixing
time
0
0 200 400 600
(c) Time (seconds)

Figure 15 (a) The mixograph (original mechanical design) used for measuring dough properties. (b) The mixing bowl is connected to a swinging arm,
which creates (c) the trace of dough properties.

Figure 17 Loaves made in testing flour quality differ in volume, height,

and crust color.

formed from a yeasted dough or from a cake batter with a

Figure 16 Scanning electron micrograph of a wheat-flour dough,
showing the strands of gluten surrounding discreet starch granules.
chemical raising agent. Cookies and crackers (Figure 3) may
or may not be leavened, but oven baking is usual.
Leavening is not needed for the wide range of pasta
(extruded from a dough of durum semolina) or noodles
(generally cut into strips from a sheet of dough made from
common wheat flour). These products (Figures 4 and 5) are
typically air-dried, permitting them to be stored for subse-
quent boiling or deep-frying (noodles) before being ready
to eat.
The range of wheat-based foods also includes the great
variety of ready-to-eat breakfast cereals, pizzas, pretzels, and
snack bars, many of which involve oven baking. Others are
formed and steam-heated by the process of extrusion.
114 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
Figure 18 Chapati bread.
Figure 19 Arabic flatbreads just taken from the oven resemble balloons
because the process is designed to separate the two outside crusts.
Figure 20 The doughnut. Heating is provided by boiling oil.
Grain Composition
Starch is the main component (up to 85%) of the mature
wheat grain. The white granules of starch are visible in the
micrograph of endosperm (Figure 11) as larger A-type and
smaller B-type granules (Figures 13 and 14). When starch is
heated in the presence of excess water, its crystalline structure is
lost in the process of gelatinization. The resulting amylose and
amylopectin macromolecules serve to bind together the other
flour components.
The major industrial uses of wheat flour involve preparing
starch by forming a dough and washing out the intact starch
granules. These are then used for a wide range of food and
nonfood purposes, the latter including the manufacture of
paper, adhesives, the sizing for textiles, hydrolysis for
sweeteners, and fermentation for biofuels.
Fats (lipids) make up only 34% of the whole grain (mainly
in the germ) and 12% of straight-run white flour. Despite
being a minor constituent, lipid plays a significant role in
dough formation and the baking process.
The grain is a significant source of micronutrients, espe-
cially minerals and B and E vitamins (but not vitamin C).
Carotenoids provide color to wheat-based products, especially
pasta from durum wheat. Much of the micronutrient content
resides in the bran and aleurone layers (Figures 10 and 11),
forming the basis for recommending the consumption of
whole-grain foods. Nevertheless, a significant amount of
aleurone cell contents is included in straight-run flour.
Another reason for the recommendation of whole-grain
foods is the contribution of dietary fiber, especially in the
form of nonstarch polysaccharides. Again, a useful proportion
of this material is present in white flour.
Protein is a very important part of the wheat grain. Protein
content typically ranges from 8% to 15%. Its importance is
indicated by the relevance of premium payments provided for
this component and the indications of utilization procedures
being based on protein content. Conventional baking use is
associated with the upper range of protein content, whereas
cake and cookie uses relate to lower protein levels.
The gluten-forming protein of white flour represents about
80% of the protein; the remaining proteins are the albumin
and globulin fractions, largely made up of proteins involved
with the synthetic functions of the grain.
Gluten is a very complex mixture of proteins about half
classed as gliadins (with a molecular weight range below
100 000 Da) and the remainder as glutenins (with a molecular
weight ranging up into the millions of Daltons). These glute-
nin proteins are large polymers of polypeptides (subunits)
joined together by disulfide bonds. The glutenin polymers,
especially the very large ones, contribute to the elasticity of
dough. To demonstrate the importance of their large size,
dough properties are lost if disulfide bonds are broken.
The industrial process of starch preparation involves the
production of gluten, which is generally dried under condi-
tions that preserve its dough-forming properties. Vital dry glu-
ten is used extensively as a bakery ingredient to increase dough
strength. It also has wide uses in many foods as a protein
extender, for example, in cheese and processed meats. In
some applications that do not call for its elastic properties,
gluten is devitalized by excess heat during drying.
Wheat and Health Issues
The many and various wheat-based foods provide a major
source of nutrition for the world. They are an important source
 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread 115

of energy, protein, essential amino acids (even if slightly defi-

cient in lysine), essential fatty acids, vitamins (with the excep-
tion of vitamin C), minerals, and fiber. For these reasons,
wheat and the wider range of cereals are generally placed at
the base of nutrition pyramids as the category of eat most.
Glycemic index and response vary between the many
wheat-based foods. Research attention has recently indicated
new genotypes and processing methods that will increase gly-
cemic index in breads and related foods.
However, certain individuals have difficulty in tolerating
wheat in their diet. These wheat intolerances take various
forms; the best categorized is celiac disease. If celiac disease is
diagnosed, the remedy is lifelong avoidance of wheat, together
with the related species rye, triticale, barley, and possibly oats.
The storage protein of all these cereals is sufficiently similar to
that of wheat gluten in amino acid composition as to cause
celiac symptoms, though not sufficiently similar as to provide
the dough-forming properties of wheat gluten.
Despite the unique requirement of wheat gluten for dough
formation, various substitutes are available for the manufac-
ture of gluten-free breads, but it has proved difficult to produce
these products to the same quality as normal bread. However,
these gluten-free foods mean that celiacs need not avoid bread-
like products completely.
The use of the word gluten to indicate both dietary intol-
erances and dough-forming properties shows it to be a misused
term, having different meanings depending on the context. Figure 21 The dripping nose of the woman at the left in this very old
Even more confusing is the term corn gluten, meaning the cartoon indicates that bakery workers have long had flour-related
protein from maize grain, although that protein has neither allergies. Reproduced with permission from Rohlich M (1978) Korn und
dough-forming properties nor toxicity for celiacs. The term Brot im Spiegel der graphischen Kunste. 7 Teil. Getreide, Mehl und Brot
glutinous rice (different spelling for this glutin) refers pri- 32: 337341.
marily to starch properties.
The possible presence of contaminants is a general risk with
wheat and other grains. These include mycotoxins and ergot. Future Prospects
However, effective screening measures at grain receival mean
that there is a very low risk that such grain will enter the human Wheat breeders have made enormous contributions to the
food chain or even animal feed. improvement of wheat for more than a century. The increases
Wheat flour is also an occupational health risk for bakers in production shown in Figure 8 are largely attributed to the
and millers who are inevitably exposed to flour on a regular contributions of breeders. Breeders will continue to provide
basis. Symptoms may include rhinitis and asthma. This occu- improvements to yield, via better tolerance to pests, better
pational problem has been recognized for centuries, as indi- adaptation to specific environments and better processing
cated by the cartoon in Figure 21. The female baker is quality as they make informed crosses, growing out the prog-
identified in the conversation as Mrs. Caquet (meaning Mrs. eny (Figure 22) and selecting for desirable traits.
Gossip). She is illustrated with nasal discharge, indicating rhi- Marker-assisted selection will continue to make wheat
nitis presumably due to flour allergy. However, the main point improvement more efficient as geneticists identify better
of the cartoon relates to the lack of hygiene in the baking trade markers and more efficient screening procedures. Improve-
at that time and the consequent health risks. The old French ments will still come from conventional breeding methods,
translates: but novel approaches to genetic modification are already per-
mitting breakthroughs that are not otherwise possible.
So you have sacked Catherine?
Wheat growers have lessons to learn from organic growers,
What else could I do Mrs Caquet, in my trade cleanliness is
essential. I couldnt keep a girl who bathed every fifteen days. She without necessarily going all the way organic. As a result,
must have been naturally very dirty as Ive never bathed in my life. agronomic strategies will improve to ensure better sustainabil-
ity and more efficient uses of agricultural inputs. The adoption
Bakers asthma is an occupational hazard that can suddenly of precision agriculture is set to increase, plus its extension to
affect bakers who have been in the trade for many years. For targeted harvesting so as to maximize the quality specifications
them, the only alternative is usually to leave baking and find a of the harvested grain.
different employment. The rhinitis and asthma are a conse- Advances in screening methods used in breeding have the
quence of immediate hypersensitivity to flour protein via potential for adaptation post-farm gate at harvest. Depending
immunoglobulin E of the nasal and bronchial tracts. In some on the economy of grain segregation, such screening methods
cases, the development of asthmatic symptoms is not immedi- can distinguish load deliveries according to suitability for uti-
ate but delayed, occurring during the night following exposure. lization and provide a basis for identity preservation of these
116 THE CEREAL GRAINS | Wheat: An Overview of the Grain That Provides Our Daily Bread
Figure 22 Breeders lines of wheat being tested in small plots for their
agronomic traits.
premium grades during storage and transport to the respective
destinations.
In these respects, wheat growers are increasingly working
on the strategy of considering the dollar value return per hect-
are, rather than grain yield, by targeting premium payments for
their grain. A further step along this approach involves a
paddock-to-plate strategy by some farming entrepreneurs,
who have milled their own grain, grown to suit quality speci-
fications, for local manufacture of food products ready to go
straight onto supermarket shelves, for example, pasta, noodles,
bread mix, and instant noodles.
Exercises for Revision
What features distinguish wheat from other cereal grains?
Review the range of growth conditions under which wheat
is grown around the world. What problems and adapta-
tions are needed to accommodate these extremes?
How do wheat and bread impact our lives and culture?
In your local region, what are the most popular uses of
wheat? Consider both food and nonfood uses.
What are the factors that affect the yield and quality of
wheat grain?
Exercises for Readers to Explore the Topic Further
The gluten protein of wheat can readily be separated from
the starch of flour by making a dough and kneading it
under a gentle stream of water. Attempt to perform this
separation to gain a better understanding of the nature of
a bread-making dough.
In your local region, find out the origins of wheat used for
the many food and nonfood products available. Is there
wheat growing nearby? Are there local milling and
manufacturing facilities?
What are the meanings of bread in your home, district,
and country?
Compare the advantages and problems of using whole-
grain flour versus white flour for wheat-based foods.
Investigate the origins and progenitors of modern wheats.
See also: Agronomy of Grain Growing: Wheat: Agronomy;
Breeding of Grains: Wheat Breeding: Exploiting and Fixing Genetic
Variation by Selection and Evaluation; Food Grains: Intolerance,
Allergy and Diseases: Cereal Allergens; Celiac Disease; The Gluten-
Free Diet; Genetics of Grains: Wheat Genetics and Genomics; Grain
Composition and Analysis: The Composition of Food Grains and
Grain-Based Products; Standardized Test Methods for Grains and
Grain-Based Products; Grain Harvest, Storage and Transport:
Wheat: Harvesting, Transport, and Storage; Grain Marketing and
Grading: Wheat: Grading and Segregation; Wheat: Marketing;
Proteins: The Protein Chemistry of Cereal Grains; The Basics: Grain:
Morphology of Internal Structure; The Cereal Grains: An Overview of
the Family of Cereal Grains Prominent in World Agriculture; Cereals:
Domestication of the Cereal Grains; Wheat Processing: Gluten and
Modified Gluten; Analysis of Dough Rheology in Breadmaking; Wheat:
Dry Milling; The Gluten Proteins of the Wheat Grain in Relation to Flour
Quality; Ultrastructure of the Wheat Grain, Flour, and Dough; Wet
Milling of Wheat; Wheat-Based Foods: Breads.
Further Reading
Anderson RP and Wieser H (2006) Medical applications of gluten-composition
knowledge. In: Wrigley CW, Bekes F, and Bushuk W (eds.) Gliadin and Glutenin.
The Unique Balance of Wheat Quality, pp. 387409. St Paul, MN: AACCI Press.
Arendt EK and Zannini E (eds.) (2013) Cereal Grains for the Food and Beverage
Industries. Oxford: Woodhead Publishing Ltd.
Baldo BA and Wrigley CW (1984) Allergies to cereals. Advances in Cereal Science and
Technology 6: 289356.
Carver B (ed.) (2009) Wheat: Science and Trade. Ames, IA: Wiley-Blackwell.
Cauvain SP (ed.) (2003) Bread Making: Improving Quality. Oxford: Woodhead
Publishing Ltd.
Cauvain SP (ed.) (2012) Bread Making: Improving Quality, 2nd edn. Oxford: Woodhead
Publishing Ltd.
Day L, Batey IL, Wrigley CW, and Augustin MA (2006) Gluten uses and food industry
needs. Trends in Food Science and Technology 17: 8290.
Faridi H and Faubion J (1995) Wheat End Uses Around the World. St Paul, MN: AACCI.
Hammed AM and Simsek S (2014) Hulled Wheats: A Review of Nutritional Properties
and Processing Methods. Cereal Chemistry 91: 97104.
Henry RJ (2014) Wheat genomics for grain quality improvement. Cereal Foods World
59: 2224.
Khan K and Shewry PR (eds.) (2009) Wheat: Chemistry and Technology, 4th edn. St
Paul, MN: AACC International.
Wrigley CW (2012) Proteins The basis of life. Teaching Science 58: 5659.
Wrigley CW and Batey IL (2010) Cereal Grains: Assessing and Managing Quality.
Cambridge: Woodhead Publishing.
 Durum Wheat: Overview
GP Kadkol and M Sissons, NSW Department of Primary Industries, Calala, NSW, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Durum wheat is a related but separate species of wheat. It is
a cereal and it belongs to the same genus as bread wheat. Its
origin is similar to bread wheat.
Durum is a tetraploid with A and B genomes represented,
which are also found in bread wheat.
Durum is adapted to more diverse environments than
bread wheat, and it performs very well in semiarid regions.
World production of durum wheat is 58% of bread wheat
production.
Durum grain is typically very hard with high protein con-
tent, amber-colored, and vitreous.
Durum has a special niche as a food crop being the main
grain used for making pasta, couscous, burghul, and several
other traditional foods.
Durum breeding objectives include higher yield, disease
resistance, higher grain protein content, increased grain
weight, low screenings, higher grain vitreosity, higher sem-
olina yield, semolina color, stronger dough characteristics,
and improved pasta-making characteristics.
Durum, similar to bread wheat, is self-pollinated, and
therefore, pedigree breeding is a common method of breed-
ing used in durum.
The main diseases of durum are Fusarium head blight,
crown rot, stem, leaf, and stripe rusts.
Management of nitrogen nutrition to the crop is an impor-
tant aspect of durum agronomy because grain protein con-
tent together with physical grain quality determines the
delivery grade and price for the farmer.
Future prospects for durum appear to be good as pasta
production has increased substantially over the previous
decade.
Learning Objectives
To learn about durum wheat as a related but separate
species of wheat and as a world food crop
To learn about the science behind the durum crop includ-
ing breeding, quality evaluation, and agronomy
Introduction
Durum wheat (Triticum durum) is a separate tetraploid species
of wheat. Durum has 28 chromosomes representing A and B
genomes compared with 42 in the hexaploid (bread) wheat
with A, B, and D genomes. The typical durum grain is very
hard, vitreous, and amber-colored with high protein content.
Durum wheat is used to produce a range of food products, such
as semolina, pasta, burghul wheat, couscous, and desserts
(Figure 1). The main durum-growing regions are the Middle
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00024-3
East, southern Europe, North Africa, the former Soviet Union,
North America (including Mexico), and India. Worldwide,
durum is grown approximately over 13 m ha with an approx-
imate annual production of 30 mt (Figure 2). Although durum
constitutes only 58% of the world wheat production, it is an
economically important crop because of its unique character-
istics and its use in making important food products such as
pasta.
Origin and Spread of Durum Wheat
It is thought that modern durum evolved from the wild tetra-
ploid, T. dicoccoides. This tetraploid, in turn, originated in the
Near East through hybridization of diploid Triticum urartu (Au
genome) and another diploid species, possibly Aegilops spel-
toides. T. dicoccoides possesses a brittle rachis and is hulled
making it difficult to harvest and hard to thresh the grain out
of the spikelets. Gradually, during domestication, these types
evolved into a tough rachis but hulled grain form, T. dicoccum,
commonly known as cultivated emmer, which is considered to
be the first cultivated tetraploid. Emmer is still hard to thresh
but can be gathered more easily compared with T. dicoccoides
because the spikes stay intact. Emmer spread from the Near
East to various European, African, and Asian regions. Archaeo-
logical evidence has shown the arrival of emmer into regions
far away from the Near East (its center of origin) into Britain, to
the West, around 3500 BCE and in the Indian subcontinent, to
the East, around 5000 BCE. Emmer is still cultivated on a small
scale in Ethiopia, India, Iran, Italy, Spain, eastern Turkey, and
the Balkans.
The process of domestication of emmer influenced by its
wide distribution and local cultural practices gradually resulted
in selection of mutations that resulted in the loss of the tight
covering of the kernel by the glumes giving rise to naked free
threshing forms. This change is thought to have occurred
approximately around 70005000 BCE. Domestication also
influenced other traits such as seed size. The first archaeologi-
cal occurrence of modern durum was recorded in Egypt during
the Greco-Roman period (332 BCE395 CE). Durum evolved
in the eastern Mediterranean region and it had established as a
major crop by 2300 BCE. Gradually, durum replaced emmer in
cultivation and spread to other parts of the world and became a
food crop of antiquity in Mediterranean Europe, several North
African and Middle Eastern countries, and the Indian subcon-
tinent and is an important ingredient of traditional foods in
these regions.
Adaptation of Durum
As a species, durum is adapted to a wide range of conditions
from temperate colder climates to warm tropical and semiarid
117
118 THE CEREAL GRAINS | Durum Wheat: Overview

Figure 1 Images from left to right: Durum wheat plants, field plots of durum wheat, durum grain, semolina (http://shop.icnussa.com.au/p/8670107/
semolinadurum-wheat-1kg-.html), pasta shapes (https://thewindroseblog.wordpress.com/2014/10/24/happy-world-pasta-day/), and couscous
(http://adashofflavour.blogspot.com.au/2010/10/moroccan-couscous-salad.html).

40000 4.00

35000 3.50
Area (1000ha) production (1000t)

30000 3.00

25000 2.50

Yield (t/ha)
20000 2.00

15000 1.50

10000 1.00

5000 0.50

0 0.00
3

1
00

00

00

00

00

00

00

01

01
/2

/2

/2

/2

/2

/2

/2

/2

/2
02

03

04

05

06

07

08

09

10
20

20

20

20

20

20

20

20

20

Figure 2 Worldwide area (brown), production (blue), and yield (blue line) statistics for durum wheat. USDA Foreign Agricultural Service http://www.
pecad.fas.usda.gov/highlights/2010/11/global%20durum/images/GlobalDurumChart.jpg.

conditions and is thus able to be grown in regions where bread
wheat may not be profitable. The majority of the world pro-
duction of durum occurs under rainfed conditions in semiarid
regions with relatively dry climate, warm days, and cool nights
during crop growth. Drought stress occurs frequently in these
regions.
Most durum varieties are spring types. In the warmer
regions of southern Europe, North Africa, and Australia, spring
durum is planted in late autumn or early winter and is
harvested in summer. In North America, spring varieties are
planted in spring and harvested in late summer. In the colder
parts of Turkey and certain other Middle Eastern countries,
winter durum varieties are planted in autumn and harvested
in summer. Winter durum is shown to be well adapted to some
areas of North America and Germany, and there is interest to
develop winter durum cropping in these regions.
Generally, durum yields are competitive with bread wheat
under favorable soil conditions. However, under low pH and

in the presence of mineral toxicities in the soil, durum yields
are typically lower than bread wheat.
Durum Breeding
Improving productivity of durum by enhancing the inherent
ability of durum varieties to produce higher yields combined
with high quality is essential for the continued viability of the
durum industry. Good varieties combining high yields and
high quality to meet the market demand have been the basis
of strong durum industries. In durum breeding, as with many
other cereals, the breeder must try to satisfy the requirements of
the entire production chain consisting of farmers, grain han-
dlers, millers, pasta makers, and consumers and is usually
assisted by experts from other fields like cereal chemistry,
agronomy, pathology, molecular biology, and biometry.
Durum breeding, historically, has been mainly conducted
by public institutions such as government departments of
agriculture and universities, while in the United States and
EU, private breeding companies are common. Durum breed-
ing commenced in EU in the early decades of the twentieth
century. Italy was a pioneer in durum breeding with the first
variety released in 1915. Breeding in the early years involved
the use of European, Near Eastern, and North African land-
races. However, these early varieties were tall, responded
poorly to nitrogen fertilizers, were prone to lodging, and were
relatively low yielding. In the 1960s, semidwarf durum germ-
plasm became available from International Maize and Wheat
Improvement Centre, Mexico (CIMMYT). Use of this germ-
plasm in durum breeding has delivered large improvements
in yield in most countries. The modern durum varieties are
shorter, more resistant to lodging, responsive to nitrogen
application, and better adapted to their target environments.
Durum Breeding Methods
Durum wheat is a self-pollinated crop and therefore a pure-line
breeding approach is used, that is, the final product is a uniform
true-breeding variety. This is achieved by several generations of
selection in the segregating material from crosses, ensuring self-
pollination at each stage. Crosses are made between carefully
chosen parent lines that have performed well in evaluation trials
or lines that carry particular traits. Many programs use not
only simple two-way crosses but also three-way crosses and
backcrosses. Most durum breeding programs use variations of
the standard pedigree breeding method (which involves making
selections under self-pollination while maintaining lineage and
identity information on the selections in each segregating gen-
eration until the breeding material reaches homozygosity), and
lines are generally bulked at F4 or F5 stage. Some programs
conduct early generation yield testing and then reselect from
high yielding bulks. The North American programs make use of
off-season nurseries in the southern hemisphere (New Zealand
or Chile) to speed up breeding. Quality testing in early stages is
limited to visual inspection of grain quality and evaluation of
protein and yellow pigment using near-infrared spectroscopy
(NIR). For further information on breeding methods, please
refer to the chapter on wheat breeding.
THE CEREAL GRAINS | Durum Wheat: Overview 119
In addition to the methods mentioned earlier, some pro-
grams also use doubled haploids (DHs) to accelerate progress
to homozygosity. DHs are completely homozygous lines gen-
erated by artificially doubling the haploid set of chromosomes
in pollen mother cells. DHs are more expensive to produce in
durum, compared with bread wheat, because of a lower success
rate. Single seed descent is a cheaper acceleration strategy and it
could typically achieve progression from F1 seed to F4 seed in a
year compared with 4 years required in the field-based conven-
tional approach.
All the previously mentioned strategies could be com-
bined with molecular marker-assisted selection to achieve
more rapid breeding progress. Molecular markers are avail-
able in durum to screen for several quality traits (e.g., grain
yellow pigment content, favorable lipoxygenase alleles, grain
cadmium content, and grain protein concentration), disease
resistance (mainly rust resistance genes), and genes involved
in determining phenology such as vernalization and photo-
period response.
Advanced evaluation of experimental lines is conducted in
replicated yield trials in several key locations in the target
regions. Detailed testing of grain, semolina, and pasta-making
quality is conducted at this stage of breeding. Testing for dis-
ease resistance is normally conducted at all stages of breeding,
but in advanced stages, the tests would include seedling resis-
tance evaluation. In some countries, varieties selected through
this process are progressed to independent variety tests where
they are evaluated against check varieties and the best new
varieties/breeding lines from other breeders.
Breeding for Yield and Agronomic Traits
Substantial improvement in the yield potential of durum has
been achieved through reducing crop stature, improving lodg-
ing resistance, and manipulating phenology to suit local grow-
ing conditions. Reduction of plant height was achieved by
incorporating the reduced height gene, Rht1, from CIMMYT,
which makes plants insensitive to the plant growth hormone,
gibberellin. The height reduction from Rht1 also resulted in
resistance to lodging and in turn made it possible to apply
higher rates of fertilizers, use irrigation (if available), and prac-
tice better agronomy to obtain higher yields.
Reducing plant height also improved the harvest index with
more of the plants resources going into grain formation. Fur-
ther improvements in productivity have come from other
improvements in plant type such as erect leaves and fine tuning
of crop development and maturity. Some breeders have pre-
ferred medium height varieties in an effort to obtain higher
yields through the tall dwarf plant type, which produces a
higher biomass. Breeding has also produced changes in repro-
ductive development phases of modern durums leading to
increases in the duration of the period between initiation of
the terminal spikelet and ear emergence resulting in improved
floret fertility and grains per spikelet.
A disadvantage of Rht1 is the reduced coleoptile length in
varieties containing this gene that makes the varieties unsuita-
ble for deep sowing. It is considered that gibberellin-sensitive
reduced height genes, such as Rht18, would result in longer
coleoptile length and provide better adaptation to deep
120 THE CEREAL GRAINS | Durum Wheat: Overview
sowing, which would allow seed placement in moist soil,
deeper down, under dry sowing conditions.
Breeding for Tolerance to Abiotic Stresses
Tolerance to moisture stress is very important considering the
majority of the durum crop is rainfed and the durum-growing
regions of the world are semiarid. In general, breeding durum
wheat for tolerance to moisture stress is currently conducted by
evaluating yield potential of the experimental lines over several
years and locations. Such evaluation will expose the experi-
mental lines to moisture stress conditions and provide oppor-
tunities to select for higher but stable yields combined with
maintenance of grain quality under drought conditions.
Tolerance to salinity has been recently researched in
Australia. Two genes, Nax-1 and Nax-2, have been incorpo-
rated into durum wheat from conventional crosses with a salt-
tolerant Triticum monococcum line. Nax-2 has been found to be
more useful for breeding because, unlike Nax-1, there was no
evidence of yield loss associated with the presence of Nax-2.
Molecular markers have been developed for these genes to
increase efficiency of selection.
Lack of tolerance to acid soils in durum germplasm has
limited the growth of durum in low-pH regions. The impact
of acid soils is through increased concentration of aluminum
in the soil solution, which is toxic to durum and limits root
growth. Recent research in Australia has achieved transfer of a
gene from bread wheat, TaALMT1, conferring tolerance to
aluminum toxicity to a popular durum cultivar, Jandaroi,
using conventional breeding methods. There is scope also for
using variation within durum germplasm as there are reports of
variation for Al tolerance comparable to bread wheat.
Breeding for Quality Traits
Durum wheat is mostly used to prepare pasta, either dried or
fresh. Other products made from durum wheat include bread
(flat and leavened), bulgur, freekeh, couscous, pastry,
pancakes, and some traditional Indian savory dishes and des-
serts. The following discussion will focus on the selection
methods and objectives in breeding for technological quality
to meet customer requirements for pasta products. In addition,
there is new interest in breeding for nutritional quality but this
will be covered elsewhere in this book.
The choice of testing procedures to select for technological
quality used in breeding programs around the world will vary
depending on the program objectives, but many breeding
targets are common across the world and will be described
here.
Key Technological Requirements
The process of selecting the many genotypes produced in
breeding programs for quality typically compares the perfor-
mance of the test line(s) with commercial (control) varieties
from the same field trial. A new test line generally needs to
match or exceed the quality characteristics of the controls. The
methods used to assess quality depend on the stage of the
breeding process that determines the amount of seed available
for testing. Generally, the earlier the generation cycle, where
seed availability is limited, the more the requirement for tests
that use only small amounts of grain/semolina with preference
for nondestructive testing. It is at these stages where there may
be several thousand lines to evaluate that the test should allow
effective screening and reduction of the population to more
manageable quantities for replicated field testing. At the later
stages of breeding, more grain is available allowing more
detailed testing.
Grain Size and Test Weight
Test weight (TW), which is affected by the size and shape of the
grain, is a widely used specification in wheat grading because it
is internationally recognized as an index of wheat soundness
and wheat milling potential. TW is lowered by grain weath-
ering, caused by rain at harvest, and grain shriveling due to
terminal drought. Shriveled, high-protein grain produces a
reduced semolina milling performance. This is because small
kernels contain a lesser percentage of endosperm, the main
economically valuable part of the grain. Generally, a high
thousand grain weight (TGW >40 g) and TW (>76 kg/hl) are
desirable and further improvements of grain size are sought.
Percentage of Hard Vitreous Kernels
Durum wheat kernels should have a translucent, vitreous
appearance to ensure good semolina yield and to avoid pasta
issues such as weak strands, poor cooking quality, and white
specks, which occur when grain vitreousness is very low
(<50%). Nonvitreous or starchy or piebald kernels tend to
produce more flour than semolina upon milling because they
are softer and the flour can give undesirable white specks in
pasta (Figure 3). Where the grain protein content is high, hard
vitreous kernels (HVK) is not usually an issue and has become
less important where millers use finer granulations. Genotypes
that produce low HVK grain compared with control varieties
having similar protein achievement across multiple environ-
ments should be selected against, but the influence of environ-
ment on protein content makes selection for HVK difficult to
achieve in practice.
% Hard vitreous kernels (HVK)
Vitreous
Piebald
Starchy
Bleached
Nonvitreous
Figure 3 Classification of grain vitreosity.

Screenings
These are undersized grains that fall through a 2 mm screen.
Grain with high percentage of such grain is undesirable for
milling because it reduces semolina yield and creates a less
uniform particle size, and this causes white spots in pasta due
to uneven hydration. Therefore, lines with a tendency to high
screenings are avoided by breeders.
Grain Protein
A key quality parameter in durum is grain protein content that
should be above 13% (11%moisture basis) to ensure about
12% protein in the semolina and pasta that is important to
obtain a good texture of cooked pasta. Dry pasta made from
high-protein semolina (12%) is physically stronger and more
elastic than pasta from lower protein semolina. Generally, as
protein content increases, cooked pasta becomes firmer and
less sticky and resists overcooking better. Negative aspects of
high protein content are lower TW and semolina yield, which
produces duller pasta. With very low levels of protein, fragile
spaghetti is produced with low firmness. High-protein durum
wheat allows spaghetti to swell when cooked, reduces cooking
loss, and allows retention of firmness with overcooking, which
is also associated with less stickiness. Typically, farmers use
nitrogen fertilizers to achieve these protein levels, but due to
the increasing cost and environmental concerns about the use
of too much fertilizer, breeders select for inherently high pro-
tein content. Significant advances have been made by breeders
with new varieties from Canada (Strongfield and CDC Verona)
and Australia (EGA Bellaroi and Jandaroi) that are inherently
high-protein achievers. Grain protein content is easily mea-
sured using NIR allowing large numbers of experimental lines
to be assessed in breeding programs.
Yellow Pigment
This is important in the market place as it contributes to the
desirable yellow color typical of pasta products that consumers
desire and provides valuable antioxidants. The yellow pigment
content in the grain is strongly inherited, which has allowed
large improvements in this trait worldwide over the last decade
from breeding. Yellow pigment can be measured using NIR or
extraction of the yellow pigment from wholemeal using sol-
vents or measurement of the semolina color with a camera to
measure the color space parameters, brightness (L*), redness
(a*), and yellowness (b*). These methods allow measurement
on small amounts of grain/semolina at very early stages in the
breeding program allowing rapid genetic gain.
Gluten Quality
Gluten strength is important in pasta manufacture and influ-
ences cooking quality (loss of solids into the cooking water)
and texture (bite, stickiness, and adhesiveness) of the cooked
product. Gluten strength is a property of the dough that has
become sought after in many markets where a higher price can
be commanded. This is because semolina from extra strong
durum varieties is thought to produce firmer pasta. The heri-
tability of gluten strength is moderate and genetic gain has
THE CEREAL GRAINS | Durum Wheat: Overview 121
been obtained over the years leading to stronger gluten types.
Gluten strength can be measured by several methods (mixo-
graph, extensograph, farinograph, glutograph, alveograph, and
gluten index) that use from 10 to 300 g of semolina that can
rank cultivars with diverse strength equally well.
Semolina Yield
A miller is mainly concerned with the milling yield or percent-
age of the grain converted to semolina of acceptable purity
with minimal flour production because semolina commands
a higher price. TW appears to be a better predictor of durum
wheat milling performance than kernel weight. Purity is
assessed as ash content (mineral content remaining when all
the organic content has been removed by combustion at very
high temperatures) and the number of brown specks per unit
area in the semolina. High mill extraction rates produce sem-
olina with higher ash (0.91.1% dry weight basis) due to
contamination by the bran and this reduces semolinas bright-
ness and yellowness. Ash content is affected by the genotype
and environment. It is desirable to avoid genotypes with a
tendency to higher ash contents. There are currently no satis-
factory small-scale predictive measures of milling performance,
and typically, a minimum of 100 g of grain has to be milled but
larger quantities (3 kg) are desirable when using a Buhler mill,
for example. The higher grain quantities required and the cost
of such milling have meant that these tests are conducted on
small elite sets of lines toward the end of the selection process,
and this necessity has made improvements in this trait difficult.
Nevertheless, a small improvement in semolina yield has been
achieved in Australia.
Pasta Quality
Evaluation of the quality of the final pasta product is the
ultimate test of acceptability of a new variety. In traditional
pasta-consuming countries, the consumer is concerned about
the aroma, color, appearance, texture, flavor, and nutritional
value of the pasta. The color of the pasta is very important
and selection of lines giving high brightness and yellowness
in uncooked and cooked pasta is desirable. This can be
achieved easily using a Minolta chroma meter or spectropho-
tometer. A high level of yellow pigment in the semolina does
not guarantee a high color in the pasta because it depends on
the activity the enzyme lipoxygenase (LOX) in the semolina
during pasta processing. LOX levels in durum wheat are
affected by genotype and environment, and assays for the
enzyme or molecular markers exist to screen genotypes for
low LOX levels.
The pasta after cooking should maintain its texture and not
become a thick, sticky mass. Mechanical texture is typically
described by a range of terms (firmness, elasticity, stickiness,
chewiness, and bulkiness) and can be measured by a sensory
panel or by objective tests with the latter typically used. Also,
cooking tests are employed to measure water absorption,
degree of swelling, and solid loss to the cooking water to assess
pasta quality.
Selection for end use quality as discussed earlier has led to
release of several varieties in many durum-growing countries
that combine high yield and high quality. Some examples are
122 THE CEREAL GRAINS | Durum Wheat: Overview
Svevo and Normanno in Italy; Strongfield, Eurostar, and
Brigade in Canada; and EGA Bellaroi, Caparoi, and Jandaroi
in Australia.
Durum Diseases and Breeding for Disease Resistance
The most important disease of durum, worldwide, is Fusarium
head blight (FHB) caused by several fungi including Fusarium
graminearum, F. culmorum, F. avenaceum, and F. sporotrichioides.
While the first two are more common causes of FHB, often,
there are more than one species present in infected crops. FHB
development is generally favored by wet conditions during
flowering and grain filling. Durum is considered more suscep-
tible to FHB than bread wheat. Infected heads are often sterile
and any seed formed is shriveled and may have a pinkish color.
FHB produces mycotoxins that make the infected grain unsui-
table for human and animal consumption. The most common
mycotoxin is deoxynivalenol for which there are legislated
maximum acceptable levels for grain trade. While there are
sources of resistance available in bread wheat and closely
related tetraploid species, resistance to FHB has been difficult
to characterize and incorporate into durum due to the influ-
ence of the environment in expression of the disease. Control
measures include adding nongrass crops in rotations and
avoiding corn that is highly susceptible to Fusarium and fungi-
cide sprays.
Fusarium spp. are also involved in causing crown rot (CR) in
durum. F. pseudograminearum and F. culmorum are the main
species involved. CR-infected plants have a honey-brown
discoloration on lower leaf sheaths and stem bases. There
could also be plant death at all growth stages. Affected heads
ripen prematurely and produce none or shriveled grain and are
known as whiteheads. Moisture stress from the flowering
stage onward promotes the disease expression and can produce
heavy yield losses. CR expression is also higher under higher
rates of nitrogen application that promotes vegetative growth
and leads to higher levels of moisture stress upon postanthesis
drought. Durum varieties are considered more susceptible to
CR than bread wheat. There is some variation for susceptibility
to CR in durum germplasm, and some progress has been
achieved in developing varieties tolerant to CR, but it is diffi-
cult to characterize the genetic effects due to large environmen-
tal effects on expression. Currently, rotation with broad-leaved
crops is the best control measure for CR.
Yellow leaf spot (tan spot) caused by Pyrenophora tritici-
repentis and Septoria leaf blotch (Mycosphaerella graminicola)
are important leaf diseases in several countries. These diseases,
being stubble-borne, are increasing in their importance as
a consequence of the increased popularity of minimum tillage.
A single recessive gene, tsn1, on chromosome 5BL controls
resistance to races 1 and 2 of tan spot. A marker is available to
screen for this gene. Resistance to races 3 and 5 is controlled by
another recessive gene, tsn2, on 3BL. The Australian varieties,
Caparoi, Jandaroi, and EGA Bellaroi, are resistant to tan spot.
While currently there are minimal yield losses in Australia
and North America from rust diseases due to good genetic
resistance, in EU countries, resistant varieties are not widely
available, and as a consequence, there is substantial reliance on
fungicides. All three rusts (stem, leaf, and stripe rust) are the
most important diseases of durum as they can produce sub-
stantial and widespread yield losses as they are airborne dis-
eases. Ensuring an adequate level of genetic resistance to all
current rust races is a major breeding objective in most breed-
ing programs. There have been significant changes of race from
time to time, and it is a significant challenge to durum breeders
and pathologists to maintain resistance in the current varieties.
For example, a new stem rust race, UG99, also known as TTKS,
has evolved in Africa overcoming two previously effective resis-
tance genes, Sr24 and Sr31. Most Australian durum varieties
carry three stem rust resistance genes, Sr8b, Sr9e, and Sr13. This
combination is expected to be effective against UG99 due to
Sr13, but it would be important to pyramid additional genes
into new varieties to improve the durability of resistance.
Resistance to leaf and stripe rust is based on adult plant resis-
tance (APR) genes. Leaf rust is normally not observed because
it develops later in the season when the weather is warm and
APR genes are effective.
Kernel discoloration (black point and red smudge) is asso-
ciated with the same fungi that cause leaf spots but are not
associated with mycotoxins. Their economic impact on durum
is from the loss of clean, vitreous appearance of the grain that
reduces the value of the grain. Durums are more susceptible to
black point than bread wheat, but there is some genetic varia-
tion in susceptibility. The main control measures are to prac-
tice good rotations to reduce fungal diseases and in irrigated
crops reduce watering after flowering.
Agronomy of Durum Production
For successful durum crop production, it is important to start
with selecting a suitable variety for the quality type targeted.
Varieties differ in their adaptation, maturity, lodging and dis-
ease resistance, and their ability to achieve the required grain
protein content and quality.
It is also important to choose a suitable paddock that does
not have a high disease inoculum load to reduce the incidence
of diseases such as crown rot. Rotation with broad-leaved
crops, such as grain legumes and canola, is very important to
reduce buildup of disease inoculum, provide opportunities for
weed control, and, in the case of grain legumes, build up soil
fertility. A good diversified crop rotation would be beneficial
economically and environmentally. It is also important to
avoid low-pH soils because of likely aluminum toxicity.
Minimum tillage or no tillage systems are gaining popular-
ity in cropping as chemical weed control is widely available.
These systems conserve moisture, improve soil structure and
soil organic matter, and reduce the cost of production. In many
regions, no tillage produces higher yields than the conven-
tional tillage systems. A disadvantage of no tillage systems is
the likelihood of disease inoculum buildup leading to higher
incidence of leaf spot diseases and crown rot. However, this
disadvantage can be overcome by adopting suitable rotations.
The timing of sowing for achieving optimum yield depends
upon location of the crop and should be determined to avoid
periods of stress including frost and terminal drought. In most
durum areas, early sowing is desirable to avoid loss of yield
potential and drought during grain development. Early sowing
allows the crop to establish and flower under favorable

conditions, and this helps reduce the impact of crown rot.
However, these benefits should be balanced against the risk
of flowering during the frost-risk period. Sowing rates are lower
for rainfed crops to conserve moisture, but in high-rainfall and
irrigated conditions, higher plant populations are targeted.
Irrigation, whether protective (strategic application of water
during spells of drought or at key growth stages) or for inten-
sive crop production, increases the yield potential of durum
crops. The modern durum varieties with short, stiff straw with-
stand irrigation and higher rates of fertilizers and produce high
yields compared with older varieties. It is important to avoid
irrigation after ear emergence to achieve high grain protein
content, HVK; to maintain good yellow pigment content; and
to reduce the incidence of kernel discoloration.
Higher rates of nitrogen nutrition are required, especially
under irrigation, to achieve the required grain protein levels. It
is possible to manage nitrogen requirements by conducting
soil tests and calculating the extra nitrogen required to achieve
the grain protein content required for the expected level of
yield. Split applications of nitrogen are more efficient at raising
grain protein content than applying all fertilizer at sowing.
With the availability of yield mapping, it is possible to apply
fertilizers more judiciously according to the observed variation
in fertility in the paddocks.
Depending upon the nutrient status of the soil, it may be
necessary to apply other nutrients such as phosphorus, sulfur,
and zinc. There are reports of both boron deficiency (Turkey)
and toxicity (Australia) in durum. Chloride deficiency and
consequent need to use chloride containing fertilizers have
been reported in the United States and it is expressed occasion-
ally in Australian crops.
Future
The future for durum as a world crop appears bright as pasta
production in the EU has increased since 2010 (Figure 4)
although global growth is coming from Asia Pacific and Middle
4.600
4.500
4.400
4.300
4.200
4.100
4.000
3.900
3.800
3.700
3.600
2003 2004 2005 2006 2007 2008 2009 2010 2011 2012
(000 tons)
Figure 4 Pasta production in the EU countries from 2003 to 2012. http://www.pasta-unafpa.org/ingstatistics2.htm.
THE CEREAL GRAINS | Durum Wheat: Overview 123
East/Africa regions with great potential for pasta consumption
in India and China. Pasta is a low-glycemic-index food and it
contains high levels of lutein, which is an antioxidant and a
vitamin A precursor. Regular intake of lutein could help pre-
vent macular degeneration.
There are significant opportunities for continued improve-
ments in productivity and quality of durum crops through
breeding and better agronomic management by developing
specific management practices to suit particular varieties.
World durum production has a potential to expand despite
the challenges from the changing climate, loss of arable land to
other uses, and competition from other crops.
There is considerable scope to improve the nutritional
quality and functional food properties of durum wheat to
expand consumer demand. Some examples of functional
food properties in durum under development are high amy-
lose content (to increase resistant starch) and soft durum (pos-
sible applications in noodles and biscuits). Development of
such durum lines could provide scope for an expanded durum
industry.
Exercises for Revision
How is durum wheat different from bread wheat?
Why do we need ongoing breeding work?
Why is grain quality important in durum wheat?
Which is the most dangerous disease of durum from a
human consumption point of view?
What is the importance of plant height in durum breeding?
Exercises for Readers to Explore the Topic Further
Check the various dry and fresh pasta products in the local
supermarket and note any differences in their appearance
and origin of the product.
124 THE CEREAL GRAINS | Durum Wheat: Overview
Check for the less popular durum foods in the local super-
market. Try finding semolina, couscous, burghul, freekeh,
and durum bread. Note whether the semolina is white or
yellow and work out the reason.
Check whether durum wheat is part of any traditional
preparations in your family.
Discuss whether you can cross durum wheat with bread
wheat.
Compare the glycemic index of pasta with other foods
including bread.
Investigate the chemical nature of yellow pigment in durum
grain and its importance for human nutrition.
Acknowledgments
The authors acknowledge assistance from various colleagues
who were very generous with information.
See also: Grain Harvest, Storage and Transport: Contaminants
of Grain; Wheat: Harvesting, Transport, and Storage; Grain
Marketing and Grading: Wheat: Marketing; The Basics: Grain and
Plant Morphology of Cereals and how characters can be used to
identify varieties; The Grain Crops: An Overview; The Cereal Grains:
Wheat: An Overview of the Grain That Provides Our Daily Bread;
Wheat-Based Foods: Pasta.
Further Reading
Abecassis J, Cuq B, Boggini G, and Namoune H (2012) Other traditional durum derived
products. In: Sissons M, Abecassis J, Marchylo B, and Carcea M (eds.) Durum
Wheat Chemistry and Technology, 2nd edn., pp. 177200. Eagan, MN: American
Association of Cereal Chemists International.
Bariana HS, Brown GN, Bansal UK, Miah H, Standen GE, and Lu M (2007) Breeding
triple rust resistant wheat cultivars for Australia using conventional and
marker-assisted selection technologies. Australian Journal of Agricultural Research
2007(58): 576587.
Blanco A, De Pace C, Porceddu E, and Mugnozza GTS (1988) Genetics and breeding of
durum wheat in Europe. In: Fabriani G and Lintas C (eds.) Durum Wheat Chemistry
and Technology, 1st edn., pp. 1746. American Association of Cereal Chemists:
MN, USA.
Bozzini A, David J, and Natoli V (2012) Origin and distribution of durum wheat genetic
diversity in the world. In: Sissons M, Abecassis J, Marchylo B, and Carcea M (eds.)
Durum Wheat Chemistry and Technology, 2nd edn., pp. 114. Eagan, MN:
American Association of Cereal Chemists International.
Clarke JM, DeAmbrogio E, Hare RA, and Roumet P (2012) Genetics and breeding of
durum wheat. In: Sissons M, Abecassis J, Marchylo B, and Carcea M (eds.) Durum
Wheat Chemistry and Technology, 2nd edn., pp. 1536. Eagan, MN: American
Association of Cereal Chemists International.
http://www.ipmcenters.org/cropprofiles/docs/NDwheat.pdf Crop profile for hard red
spring and durum wheats in North Dakota.
Engel RE, Bruebaker L, and Emborg TJ (2001) A chloride deficient leaf spot of durum
wheat. Soil Science Society of America Journal 65: 14481454.
Grant CA, Di Fonzo N, and Pisante M (2012) Agronomy of durum wheat production.
In: Sissons M, Abecassis J, Marchylo B, and Carcea M (eds.) Durum Wheat
Chemistry and Technology, 2nd edn., pp. 3755. Eagan, MN: American Association
of Cereal Chemists International.
Isidro J, Alvaro F, Royo C, Villegas D, Miralles DJ, and Garca del Moral LF (2011)
Changes in duration of developmental phases of durum wheat caused by breeding
in Spain and Italy during the 20th century and its impact on yield. Annals of Botany
107: 13551366.
Joppa LR and Williams ND (1988) Genetics and breeding of durum wheat in the United
States. In: Fabriani G and Lintas C (eds.) Durum Wheat Chemistry and Technology,
1st edn., pp. 4768. Eagan, MN: American Association of Cereal Chemists.
Kimber G and Sears ER (1987) Evolution in the genus Triticum and the origin of
cultivated wheat. In: Heyne EG (ed.) Wheat and Wheat Improvement, pp. 154164.
Madison, WI: American Society of Agronomy.
http://en.magazinebbm.com/?p235 The future of pasta is in less developed
countries.
Martia J and Slafer GA (2014) Bread and durum wheat yields under a wide range of
environmental conditions. Field Crops Research 156: 258271.
Matthews P, McCaffery D, and Jenkins L (2014) Winter crop variety sowing guide 2014.
NSW Department of Primary Industries. http://www.dpi.nsw.gov.au/agriculture/
broadacre/guides/winter-crop-variety-sowing-guide.
Morris CE and Sands DC (2006) The breeders dilemma Yield or nutrition? Nature
Biotechnology 24(9): 10781080.
DePauw R and OBrien L (2014) Wheat Breeding: Exploiting and Fixing Genetic
Variation by Selection and Evaluation. In: Wrigley C, Seetharaman K, Corke H, and
Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn, vol. 4, pp. 279286.
Elsevier Ltd.
Park RF (2014) Wheat: Biotrophic Pathogen Resistance. In: Wrigley C, Seetharaman K,
Corke H, and Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn, vol. 4,
pp. 264272. Elsevier Ltd.
Periyannan SK, Qamar ZU, Bansal UK, and Bariana HS (2014) Development and
validation of molecular markers linked with stem rust resistance gene Sr13 in durum
wheat. Crop & Pasture Science 65: 7479.
Royo C, Nachit M, Di Fonzo N, Araus JL, Pfeiffer M, and Slafer JA (2005) Durum Wheat
Breeding Current Approaches and Future Strategies. volume 1, New York, NY:
Food Products Press.
Sissons MJ (2004) Pasta. In: Wrigley C, Corke H, and Walker C (eds.) Encyclopedia of
Grain Science, pp. 409418. Australia: Elsevier.
Sissons MJ (2014) Pasta. In: Wrigley C, Seetharaman K, Corke H, and Faubion J (eds.)
Encyclopedia of Food Grains, 2nd edn, vol. 3, pp. 7990. Elsevier Ltd.
Sissons M, Abecassis J, Marchylo B, and Cubadda R (2012) Methods used to assess
and predict quality of durum wheat, semolina and pasta. In: Sissons M, Abecassis J,
Marchylo B, and Carcea M (eds.) Durum Wheat Chemistry and Technology,
2nd edition, pp. 213234. Eagan, MN: American Association of Cereal Chemists
International.
Torrey G Lutein May Decrease Your Risk of Macular Degeneration. American Macular
Degeneration Foundation, https://www.macular.org/lutein.
Troccoli A, Borrelli GM, De Vita P, Fares C, and Di Fonzo N (2000) Durum wheat
quality: A multidisciplinary concept. Journal of Cereal Science 32: 99113.
Wallwork H (2000) Cereal Leaf and Stem Diseases, GRDC and SARDI, ISBN 1 875477
86 1.
Wallwork H (2000) Cereal Root and Crown Diseases, GRDC and SARDI, ISBN 1 875477
74 8.
http://herbarium.usu.edu/webmanual/info2.asp?
nameTriticum_durum&typetreatment Wheat taxonomy.
Wrigley C (2016) Wheat - An Overview of the Grain that provides Our Daily Bread. In:
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 Rice: Overview
BO Juliano, Philippine Rice Research Institute Los Banos, Laguna, Philippines
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Rice is the most important crop in the world in terms of
total developing world production and the number of con-
sumers dependent on it as their staple.
Semidwarf and hybrid rice, together with tolerance/
resistance to abiotic and biotic stresses, have increased rice
production.
Rice is hypoallergenic with protein of good quality, and its
net protein utilization (NPU) is not affected by cooking.
Its wide range of apparent amylose content (AC) has con-
tributed to wide differences in eating quality preferences for
table rice and for rice products within the various regions
of Asia.
Differences in starch gelatinization temperature and gel and
Rapid Visco Analyser consistency are due to differences in
amylopectin properties, where amylose content is similar.
Glycemic index decreases with increase in AC.
g-Oryzanols and tocols in unsaponifiable matter of unde-
fatted brown rice and bran have hypocholesterolemic
properties.
Learning Objectives
To appreciate the wide adaptability of the rice plant and the
unique properties of the rice grain
To be able to relate structure and composition of rice to its
nutritional properties
To understand the diversity in rice texture and the causes of
textural differences
Commercial Importance, Global Distribution,
and Varieties
Rice is the most important crop in the world in terms of total
developing world production (480
106 tonnes of rough rice
in 2012) and the number of consumers (3.5 billion) depen-
dent on it as their staple food. It is widely grown in over 100
countries in every continent (except Antarctica), from 53 N to
40 S and from sea level to an altitude of 3 km. The total
production in 2012 was 697
106 tonnes of rough rice. Asia
accounts for 149
106 of the 158
106 ha world area that is
used for rice cultivation. The mean rough rice yield was 4.40 -
tonnes ha1 in 2012. About 90% of the worlds rice is grown
and consumed in Asia, in 200 million rice farms most of which
are smaller than 1 ha. The major rice producers in 2012 were
China, India, Indonesia, Bangladesh, Vietnam, and Thailand.
Rice can grow in both flooded (anaerobic soil) and upland
(aerobic) conditions. The plant has air channels that connect
to its roots. In terms of water regime, 54% of the total rice area
was irrigated in 1991, 25% rainfed lowland, 13% upland, and
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00015-2
9% flood-prone wetland, corresponding to a total rough rice
production of 76% from irrigated, 16% from rainfed lowland,
4% from upland, and 4% from flood-prone wetland. Since the
rice-growing area is shrinking, rice production must keep up
with the annual increase in population in tropical Asia through
increased yield.
Rice is mainly the species Oryza sativa L., plus Oryza
glaberrima L., grown in West Africa. There are estimated to be
about 100 000 rice varieties; only a small proportion is actually
widely cultivated. They vary in grain weight, size and shape,
degree of dormancy, longevity, and seedling vigor, and some
have red to purple-black pigments. About 75% of Asias rice
area is planted with varieties of improved semidwarf plant type
with erect leaves. The newer, improved varieties have a similar
yield potential to the first variety (IR8) but have a better resis-
tance or tolerance to biotic and abiotic stresses. Although
essentially self-pollinated, three-line cytoplasmic male sterile
hybrids and two-line thermogenic male sterile hybrids have
been successfully produced and grown first in China with
increased yields. Complete sequencing of the rice genome in
2002 has accelerated the use of molecular markers in breeding
for yield, stress tolerance, and, lately, grain quality.
Structure of the Rice Grain
The rice grain (rough rice or paddy) consists of an outer pro-
tective covering, the hull (husk), and the edible rice caryopsis
or fruit (brown, cargo, dehulled, or dehusked rice) (Figure 1).
Brown rice consists of the outer layers of pericarp, seed coat,
nucellus, and germ or embryo, which are the maternal tissues,
and the endosperm. The endosperm consists of the aleurone
layer and the starchy or inner endosperm. Pigment is confined
to the pericarp, but there is a varietal difference in the extent of
retention of pigment with the degree of milling. The aleurone
layer encloses the embryo.
The inedible hull constitutes 1628% (mean 20%) of
rough rice weight. Brown rice consists of 12% pericarp,
46% aleurone plus nucellus and seed coat, 1% embryo, 2%
scutellum, and 9091% endosperm. The aleurone and embryo
cells are rich in lipid bodies (spherosomes, 0.21.5 mm) and in
protein bodies (aleurone grains containing inclusions of phytic
acid bodies or globoids (13 mm)).
The endosperm cells are thin-walled and packed with amy-
loplasts containing polyhedral compound starch granules about
39 mm in size. Protein occurs mainly in the form of large
(12 mm) and small (0.50.8 mm) spherical protein bodies
and crystalline protein bodies (24 mm). Spherical protein bod-
ies (PB I) are rich in prolamin (alcohol-soluble protein), and
crystalline protein bodies (PB II) are rich in glutelin (alkali-
soluble protein). High night ripening temperature increases PB
II/PB I ratio. Spherosomes are present in the subaleurone or the
two outermost cell layers of rice endosperm.
125
126 THE CEREAL GRAINS | Rice: Overview
Lemma
Hull
Palea
Inner
endosperm
Starchy
endosperm
Subaleurone
layer
DORSAL
Sterile lemmae
Figure 1 Longitudinal section of the rice grain. Reproduced with permission from Juliano BO (1984) Rice starch: Production, properties and uses. In:
Whistler RL, BeMiller JN, and Paschall EF (eds.). Starch Chemistry and Technology, 2nd edn., p. 509. Orlando, FL: Academic Press.
Grading, Handling, and Storage
There is no international standard for milled rice size and
shape. The International Rice Research Institute (IRRI) uses
the following scale for size: extra long, >7.50 mm; long,
6.617.50 mm; medium, 5.516.60 mm; and short,
<5.50 mm. For grain shape based on the length/width ratio,
the following scale is used: slender, > 3.0; medium, 2.13.0;
bold, 1.12.0; and round,  1.0. Grades are based on grain size
and shape, degree of milling, percentage of head or whole-
grain milled rice, immature grains, damages (discolored) and
heat-damaged grains (chalky grains, red grains, and red-
streaked grains), aroma, and organic extraneous matter and
inorganic extraneous matter.
An important property of harvested grain is its moisture
content: 14% on a wet-weight basis is considered a safe storage
value; grains become susceptible to fissuring from moisture
adsorption stress below critical moisture contents of 1216%,
depending on the variety. Aging of sun-/oven-dried rough rice
for up to 2 months after harvest at ambient temperature
>15  C improves milling yields and makes milled rice expand
more during cooking, with less dissolve solids, and become
more flaky. Heating during grain drying accelerates aging.
About 20% of rice is consumed as parboiled rice. Parboiling
consists of boiling or steaming steeped aged rough rice until
the hull starts to open and then cooling and drying the gelati-
nized grain. Diffusion of bran B vitamins into the endosperm
occurs during parboiling but with some heat decomposition.
Mycotoxins are a potential problem in parboiled rice.
Milling involves dehulling, followed by removal of the
outer 710% of the brown rice, either by friction or by abra-
sion, removing most of the pericarp, seed coat, nucellus,
Awn
Pericarp
Seedcoat
Nucellus
Aleurone layer
VENTRAL
Scutellum
Epiblast
Plumule Embryo
Radicle
Rachilla
aleurone layer, and the germ. In the Engelberg mills, hull and
bran are removed together in one step with high grain break-
age. Milling is done in several steps in modern cone mills with
tempering in between to minimize breakage. Many modern
mills have shifted to milling at >14% moisture to minimize
breakage and to moisture mist treatment (through hollow
shaft) during milling to soften the bran and improve surface
gloss.
Only about 45% of the worlds rice production enters the
international trade. The major exporters in 2012 were Thai-
land, India, Vietnam, the United States, and Pakistan. The
major importers in 2012 were Nigeria, China, Iran, the Philip-
pines, Iraq, Saudi Arabia, Ivory Coast, and Malaysia.
Chemical and Nutritional Composition
Rice has one of the lowest protein contents (7%) among the
cereals. The bran layers and embryo are richer in nonstarch
constituents than the milled (white) rice (Table 1). The major
nutritional advantage of brown rice over milled rice is its
higher content of B vitamins. Although higher in minerals,
bran phytic acid forms complexes with minerals and proteins,
reducing their bioavailability. Thus, the available iron (mg per
meal) is similar in brown and milled rice meals, but probably
higher in brown rice for zinc (mg per meal) than for milled
rice. The energy content of brown rice and bran is higher than
that in milled rice, owing to the higher fat content. Rice has no
vitamin A, C, or D.
Although cereal proteins are deficient in lysine, rice protein
has one of the highest lysine contents among them, corre-
sponding to an amino acid score of 67% in milled rice based
 THE CEREAL GRAINS | Rice: Overview 127

Table 1 Comparison of nutrient composition of brown rice, milled rice, and rice bran

Amounts (per 100 g)

Property Brown rice Milled rice Rice bran

Moisture (g) 14.0 14.0 14.0

Energy content (kJ) 14801610 14601560 16701990
Energy content (kcal) 355385 349373 399476
Crude protein (g) 7.18.3 6.37.1 11.314.9
Crude fat (g) 1.62.8 0.30.6 15.019.7
Crude fiber (g) 0.61.0 0.20.5 7.011.4
Crude ash (g) 1.01.5 0.30.8 6.69.9
Available carbohydrates (g) 7387 7789 3462
Total dietary fiber (g) 2.94.4 0.72.7 2429
Water-insoluble fiber (g) 2.0 0.5 1527
Sugars (g) 0.81.9 0.10.5 5.56.9
Thiamin (mg) 0.40.6 0.070.17 1.22.5
Riboflavin (mg) 0.040.14 0.020.06 0.180.43
Niacin (mg) 3.56.2 1.32.5 2750
Pantothenic acid (mg) 1.41.6 0.81.3 2060
Vitamin B6 (mg) 0.50.7 0.10.4 3.7
Folate (mg) 1620 49 40140
Vitamin E, a-tocopherol (mg) 0.82.5 0.10.3 315
Calcium (mg) 1050 1030 30120
Phosphorus (mg) 0.170.43 0.080.15 1.12.5
Phytic acid P (mg) 0.130.27 0.020.07 0.92.2
Iron (mg) 1.45.2 0.30.8 8.643
Zinc (mg) 1.92.8 0.82.3 4.326

Juliano BO (2007) Rice Chemistry and Quality. Munoz, Nueva Ecija: Philippines Rice Research Institute; Champagne ET, Wood DF, Juliano BO, and Bechtel DB (2004) The rice grain
and its gross composition. In: Champagne ET (ed.). Rice Chemistry and Technology, 3rd edn., pp 77108. St. Paul, MN: American Association of Cereal Chemists; United States
Department of Agriculture (2012) National Nutrient Database for Standard Reference, Release 25. http://www.ars.usda.gov/ba/bhnrc/ndl.

Table 2 Amino acid profile and energy and nitrogen balance in five growing rats of raw brown rice, milled rice, and rice bran

Property Brown rice Milled rice Rice bran

Arginine (gram per 16 g N) 7.2 7.9 7.5

Histidine (gram per 16 g N) 2.4 2.2 2.5
Isoleucine (gram per 16 g N) 4.0 4.1 4.0
Leucine (gram per 16 g N) 7.9 7.8 7.3
Lysine (gram per 16 g N) 3.6 3.4 4.6
Methionine (gram per 16 g N) 2.1 2.2 2.2
Methionine cystine (gram per 16 g N) 3.3 4.2 4.4
Phenylalanine (gram per 16 g N) 4.9 5.1 4.5
Phenylalanine tyrosine (gram per 16 g N) 8.5 8.3 7.5
Threonine (gram per 16 g N) 3.5 3.4 4.0
Tryptophan (gram per 16 g N) 1.2 1.1 0.8
Valine (gram per 16 g N) 5.6 5.8 6.3
Digestible energy (% of total) 94.3b 96.6a 67.4c
True digestibility (TD, % of diet N) 96.9b 98.4a 78.8c
Biological value (% of digested N) 68.9b 67.5b 86.6a
Net protein utilization (% of diet N) 66.7b 66.4b 68.3a
Amino acid score (%) 71lys 67lys 90lys
Amino acid score
TD (%) 68lys 66lys 71lys

Means in the same line followed by a common letter are not significantly different at the 5% level by Duncans (1955) multiple range test. lys lysine is the first-limiting amino acid.
United States Department of Agriculture (2012) National Nutrient Database for Standard. Reference, Release 25. http://www.ars.usda.gov.ba/bhnrc/ndl; Eggum BO, Juliano BO, and
Maningat CC (1982) Protein and energy utilization of rice milling fractions by rats. Qualitas Plantarum-Plant Foods for Human Nutrition 31: 371376.

on the US Food and Nutrition Board amino acid requirement
pattern for the 13-year-old child of 5.1% lysine as 100%
(Table 2). The solubility fractions of protein are about 15%
of albuminglobulin (water- and salt-soluble), 20% prolamin
(PB I, alcohol-soluble), and 65% glutelin (PB II, alkali-soluble)
in milled rice. Bran proteins are 6698% albumins. Prolamin
is poor in lysine but rich in sulfur amino acids. The high lysine
content of rice protein is due to the low prolamin content.
The energy digestibility is higher in milled rice than in
brown rice owing to the lower dietary fiber and phytic acid
128 THE CEREAL GRAINS | Rice: Overview
levels, as verified by the poor energy digestibility of rice bran
(Table 2). The true digestibility (TD) of milled rice protein is
also higher than that of brown rice, but the biological value
(BV) is lower, resulting in a similar NPU. Bran protein has a
lower TD but a higher BV than brown and milled rice proteins.
The amino acid score corrected for TD in rats proposed by the
Food and Agriculture Organization as a protein quality index
shows similar values to NPU for the rice proteins. Black or
purple rice has a lower brown rice NPU (72%) and higher
phenolics level (0.6%, anthocyanin) than red rice (NPU;
83% and 0.2% phenolics, proanthocyanidins) and
nonpigmented rice (NPU 97% and 0.02% phenolics), but
their milled rice have identical NPUs. Rice complements
legumes (deficient in sulfur amino acids) in amino acid com-
position for human diets.
Cooking and parboiling reduces the TD in growing rats by
515%, with a corresponding increase in BV but little change
in NPU. However, lysine digestibility remains close to 100%,
and cystine digestibility drops to about 82%. The fraction that
remains in the feces as fecal protein particles represents the
lipid-rich core of large PB I, with < 1% lysine in the protein but
rich in cystine. PB II is readily digested.
Starch varies in apparent AC (as determined by iodine
colorimetry in acetate buffer): waxy, 02%; very low, 210%;
low, 1020%, intermediate, 2025%; and high, >25%, all on a
milled rice dry-weight basis since the 1950s. Actual AC values
by differential scanning calorimetry (by melting or crystalliza-
tion enthalpy of amyloselysolecithin complex) are slightly
lower and closer to AC values with iodine colorimetry in
ammonium buffer pH 9 of low, 1017%; intermediate,
1722%; and high, >22%. The glycemic index of cooked
brown rice tends to be lower than that of cooked milled rice
owing to the higher amounts of phytic acid and fiber in brown
rice. However, the lowering in glycemic index from milled rice
to brown rice is highest in waxy and low-AC rice. Among
cooked milled rice, the glycemic index decreases with increas-
ing AC regardless of the cooking method. Processing, including
parboiling, tends to reduce the glycemic index. In vivo resistant
starch in an ileostomate (large intestine surgically removed) is
5% and may be not as discriminating as the glycemic index
in humans. Short-term satiety may also be influenced by AC.
The use of rice bran and brown rice in cereal products has
increased in recent years owing to the US Food and Drug
Administrations recent approval of brown rice as a whole
grain and the hypocholesterolemic effect, in humans, of the
factors in the high (up to 4.4%) unsaponifiable fraction of
its oil, such as g-oryzanol, (cycloartenol, 24-methylene-
cycloartanol and campestrol esters of ferulic acid), tocopherols,
and tocotrienols, an analog of tocopherol (vitamin E). Defatted
bran has no hypocholesterolemic activity, unlike in oats,
where the active principle is soluble b-glucan. Inactivation of
antinutritional factors trypsin inhibitor, oryzacystatin, and
hemagglutininlectin and lipase and lipoxygenase that are
concentrated in the rice bran by heat treatment and extrusion
cooking improves the shelf life of the bran and its nutritional
value to poultry. By contrast, the antinutritional factor phytic
acid is heat-stable. Phytic acid in rice bran and brown rice has
been reported to have some medicinal activity in preventing
some types of cancer. Phytic acid content of rice bran is highest
among cereal brans (38% phytic acid).
Essential fatty acid content of rice oil is 33.4% linoleic acid
and 1.6% linolenic acid. Levels of oryzanol, tocopherols,
tocotrienols, and unsaponifiable matter differ among crude
oils, and oryzanol, tocopherols, and tocotrienols may be
reduced by up to 90% by conventional refining and deodoriz-
ing. Contents of these antioxidants are also lower in Philippine
rice than in US rice.
Breeding efforts to improve the nutritional value of rice
grain include a higher micronutrient density (higher iron,
zinc, and vitamin A), low phytic acid (high inorganic phos-
phate) content, low prolamin content, the absence of
lipoxygenase-3 activity and allergenic globulin, and transgenic
rice endosperm with vitamin A, with the soybean glycinin gene
and with the soybean ferritin gene.
Processing and Food Uses
The mean worldwide per capita consumption of milled rice
was 53 kg per year in 2009. It was 77 kg in Asia, 31 kg in South
America, 21 kg in Africa, 14 kg in Oceania, and <10 kg in
Europe and North America. Per capita, rice consumption was
highest in Southeast Asia at 129 kg per year, followed by South
Asia at 71 kg per year. More than 3.5 billion people depended
on rice for more than 20% of their daily energy in 2009. Rice
provides more than 50% of energy supply for about 520 mil-
lion people in Asia, most of them poor to very poor.
There are various ways of cooking milled rice. In tropical
Asia, they are prewashed to remove dirt, which results in losses
of B vitamins and fat. Presoaking for 30 min reduces cooking
time (particularly of parboiled and brown rice). Grain cracking
during soaking is minimized by adjusting milled rice moisture
content to 15% by high-pressure hydration to minimize crack-
ing. Rice may be cooked in the amount of water that it will
absorb or boiled in excess water and the cooking water dis-
carded. A steaming step is used in Indonesia and also for waxy
rice. Oil or ghee may be added in the Middle East, and this
reduces surface stickiness. Enriched rice premixes containing
iron and B vitamins resistant to washing have been developed
for rice, but have not been popular because of additional
expense; the enriched rice premix can be readily distinguished
from ordinary rice.
Rice is hypoallergenic. Its AC correlates positively with
water absorption and volume expansion during cooking and
with hardness of boiled rice. South Asian consumers prefer
high-AC intermediate-high gelatinization temperature (GT
>70  C) rice; Southeast Asians prefer intermediate-AC,
intermediate-GT rice. Japonica (temperate) short-grain rice pre-
ferred in Japan, Korea, Taiwan, and northern China have
mainly low AC and low GT (70  C), with some having an
intermediate AC in Europe. Only in Laos and north and north-
east Thailand is steamed waxy (glutinous) rice consumed as a
staple. The low GT and intermediatehigh GT result from the
differences in proportion of debranched amylopectin with
chain lengths below 24 glucose units: Low GT has more of
the shorter chains. The higher longer chains increase not only
GT but also the staling of cooked starch. Among waxy and low-
AC rice, low-GT rice is a softer cooked rice than high-GT rice
(intermediate GT is rare at low AC), but among intermediate-
and high-AC rice, intermediate-GT rice is a softer freshly

cooked rice than low-GT rice. High-AC rice with high content
of long-chain amylopectin (blue staining with iodine) have
hard gel consistency, as long-chain amylopectin behaves like
amylose. Native long-chain amylopectin crystallizes from
water-saturated 1-butanol only after debranching with
isoamylase.
Various rice products are prepared for which specific AC
types are preferred. Freshly and well-milled rice is preferred for
rice products to prolong shelf life by minimizing fat rancidity in
the stored products. Parboiled rice are preferably aged high- and
intermediate-AC, whereas extruded and flat noodles use mainly
aged, high-AC, low-GT rice. Rice with a low GT of starch is
preferred in rice puddings, breads and cakes, and beer adjuncts.
Waxy and low-AC rice are preferred for rice wines (for a higher
ethanol yield) and in frozen sauces, desserts, snacks, and sweets,
because of their slow staling rate. Rice crackers are prepared
from waxy and nonwaxy rice. Parboiled rice is preferred over
raw rice for idli (pudding) and dosai (cake) with rice-black gram
usually fermented at a 3:1 weight ratio. The thermophysical
properties of starch such as glass transition temperature, GT,
enthalpy of raw starch, staled amylopectin melting
(4560  C), amyloselipid complex I (<100  C) and II
(>100  C) melting, and staled amylose melting (>130  C) of
gelatinized starch affect the properties of rice products in addi-
tion to the amyloseamylopectin ratio and protein content.
Specialty rice include Italian Arborio rice for making risotto,
waxy rice (sweet rice with an opaque grain), Thai Jasmine rice
(aromatic low-AC long-grain), and Punjab Basmati (aromatic
intermediate-AC long-grain that elongates when presoaked rice
is cooked). 2-Acetyl-1-pyrroline is the first major aroma prin-
ciple identified in raw and cooked aromatic rice, and others are
being characterized. Endosperm 2-acetyl-1-pyrroline bound
to starch requires a higher extraction temperature than the
free form.
Exercises for Revision
While total maize production exceeds that of rice, why is
rice a more important food crop than maize?
Does waxy or glutinous rice contain gluten?
Why was the approval by the US Food and Drug Adminis-
tration of brown rice as a whole grain delayed as compared
with the other cereal grains?
The N content of rice glutelin is 16.8%. What Kjeldahl N
factor would one use for rice glutelin? Note: The factor 6.25
is used for most proteins containing 16.0% N.
Why is it harder to manipulate the composition of a whole
grain such as rice than that of a flour such as wheat?
Exercises for Readers to Explore the Topic Further
What factors would influence satiety index of cooked
milled and brown rice?
How would a low-phytic acid mutant affect iron absorption
from brown rice?
Speculate on how global warming would affect rice grain
properties.
THE CEREAL GRAINS | Rice: Overview 129
How may mutants low in lipoxygenase and lipase improve
shelf life of rice?
How does amylose differ from long-chain amylopectin in
properties?
Speculate on the binding of 2-acetyl-1-pyrroline with
starch.
See also: Barley, Rice and Maize Processing: Oil from Rice and
Maize; Rice Processing: Beyond the Farm Gate; Beverages from
Grains: Beverages: Asian Alcoholic Beverages; Breeding of Grains:
Rice: Breeding; Carbohydrates: Glycemic Index, Glycemic Load, and
Their Health Benefits; Resistant Starch and Health; Starch: Analysis of
Quality; Starch: Starch Architecture and Structure; Food Grains:
Intolerance, Allergy and Diseases: Nutrition: Beriberi, A
Deficiency Related to Grains; Food-quality Testing: Rice: Eating
Quality; Genetics of Grains: Rice: Genetics; Non-wheat Foods:
Rice: Chinese Food Uses; Scientific Organizations Related to
Grains: Research Organizations of the World: CGIAR; The Cereal
Grains: African Rice (Oryza glaberrima): A Brief History and Its
Growing Importance in Current Rice Breeding Efforts; An Overview of
the Family of Cereal Grains Prominent in World Agriculture.
Further Reading
Atkinson FS, Foster-Powell K, and Brand-Miller JC (2008) International tables of
glycemic index and glycemic load values. Diabetes Care 31: 22812283.
Biliaderis CG, Tonogai JR, Perez CM, and Juliano BO (1993) Thermophysical
properties of milled rice starch as influenced by variety and parboiling method.
Cereal Chemistry 79: 512516.
Champagne ET, Wood DF, Juliano BO, and Bechtel DB (2004) The rice grain and its
gross composition. In: Champagne ET (ed.) Rice: Chemistry and Technology,
3rd edn., pp. 77108. St Paul, MN: American Association of Cereal Chemists.
Eggum BO, Juliano BO, and Maningat CC (1982) Protein and energy utilization of rice
milling fractions by rats. Qualitas Plantarum-Plant Foods for Human Nutrition
31: 371376.
Food and Nutrition Board Institute of Medicine (2002) Dietary References Intakes for
Energy, Carbohydrates, Fiber, Fat, Fatty Acids, Cholesterol, Protein and Amino
Acids (Macronutrients). Washington, DC: National Academy Press.http://www.nap.
edu/opcabook/html/.
Hanashiro I, Matsugasako J, Egashira T, and Takeda Y (2005) Structural
characterization of long unit-chains of amylopectin. Journal of Applied Glycoscience
52: 233237.
International Rice Research Institute, 2013. World Rice Statistics. irri.org/world-rice-
statistics.
Juliano BO (1985) Rice starch: Production, properties and uses. In: Whistler RL,
BeMiller JN, and Paschall EF (eds.) Starch Chemistry and Technology, 2nd edn.,
p. 509. Orlando, FL: Academic Press.
Juliano BO (2004) Rice/overview. In: Wrigley C, Corke H, and Walker CE (eds.).
Encyclopedia of Grain Science, vol. 3, pp. 4148. Oxford, UK: Academic Press/
Elsevier.
Juliano BO (2007) Rice Chemistry and Quality. Munoz, Nueva Ecija: Philippine Rice
Research Institute.
Juliano BO, Tuano APP, Monteroso DN, et al. (2012) Replacement of acetate by
ammonium buffer to determine apparent amylose content of milled rice. Cereal
Foods World 57: 1419.
Tuano APP, Xu Z, Castillo MB, et al. (2011) Content of tocols, g-oryzanol and total
phenolics and grain quality of brown rice and milled rice applied with pesticides and
organic and inorganic nitrogen fertilizer. Philippine Agricultural Scientist
94: 211216.
United States Department of Agriculture, 2012. National Nutrient Database for Standard
Reference, Release 25.http://www.ars.usda.gov/ba/bhnrc/ndl.
Yoshihashi T, Huong NTT, Surojanametakul V, Tungtrakul P, and Varanyanond W
(2005) Effect of storage conditions on 2-acetyl-1-pyrroline content in aromatic rice
variety, KhaoDawk Mali 105. Journal of Food Science 70: S34S37.
 Wildrice, Zizania: Overview
EA Oelke, University of Minnesota, St. Paul, MN, USA
RA Porter, University of Minnesota-NCROC, Grand Rapids, MN, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Wildrice (wild rice), Zizania, belongs to the grass family, is a
cereal grain, and has four species, three naturally occurring
in North America and one in Asia.
The grain from wildrice growing in lakes and rivers in North
America has been hand harvested, using a canoe, and pro-
cessed by Native Americans for thousands of years as a
carbohydrate food.
Most cereal grains have been under cultivation and domes-
tication for centuries, but for wildrice, this first began in the
early 1950s in Minnesota.
Improved cultivars have been developed for better yields
than natural plants and adapted for harvesting with large
combines.
Wildrice cultivation is similar to rice, Oryza, in diked fields
(paddies) to allow flooding and draining.
Today, most of the cultivation is done in Minnesota and
California, the United States.
Yields per hectare of cultivated wildrice are about ten times
that from hand harvested natural stands, but still well
below that of rice.
Processing of the cultivated grain follows similar steps used
by Native Americans of parching (roasting and drying) and
hulling, however using modern equipment in inspected
processing facilities.
Processed wildrice is a unique, specialty cereal grain with a
distinct flavor and has a good amount of protein, similar to
wheat, and twice the dietary fiber of brown rice.
The processed grain (kernel) without hulls is very versatile
and is often featured in soups, salads, breads, side dishes,
entrees, and even desserts.
In Minnesota, any packaged processed wildrice sold in the
state must be labeled as to where and how it is grown.
Learning Objective
To learn about this native North American naturally growing
aquatic grass plant, wildrice (Zizania), which is in the very
early stage of cultivation/domestication as a cereal grain.
Introduction
Wildrice (Zizania sp.) is in its infancy in cultivation and domes-
tication compared to other cereals such as rice (Oryza sativa L.).
Rice has been under this process for thousands of years. In the
North American continent, the Native Americans did some
expansion of natural stands by hand seeding new lakes and
areas of lakes with sparse stands. But it was not until 1950 that
a 0.5 ha paddy in Minnesota was specifically constructed to
grow wildrice. Subsequently, more hectares were developed,
130 Encyclopedia of Food Grains, Second Edition
and some breeding efforts were initiated in the early 1960s
with a full-scale breeding program started in 1972 by the
University of Minnesota. Today, wildrice is cultivated mainly
in Minnesota and California in the United States and outside of
the United States on a few hectares in Hungary. The Canadians
have expanded lake production by seeding new lakes in several
provinces. In Canada, the natural stands as well as newly seeded
lakes are harvested by airboats with catchers attached to the
front. There are three species of wildrice that grow naturally in
the North American continent: Zizania aquatica L., Zizania palus-
tris L., and Zizania texana A.S. Hitchc. Z. aquatica grows along the
eastern seacoast and has thin kernels. Z. palustris grows around
the Great Lakes region and has large seeds. It was gathered by
Native Americans for thousands of years as a food source. Both
of these species are annual aquatic grasses. Z. texana is a peren-
nial with small seeds and grows only in a very small area in
Texas. Z. latifolia Turcz. grows in Asia, and the fungal infected
base of the stem is harvested as a vegetable, Makomo-taki.
Wildrice is a recently cultivated and partially domesticated
cereal grain; thus, the purpose of this article is to acquaint the
reader with this new crop. Wildrice is cultivated using similar
practices as is used for lowland rice, in that it only grows in
flooded soils. Harvesting the newly developed cultivars is done
with combines similar to ones used for rice in the United
States. Processing is different, however, giving wildrice its
unique roasted flavor. The nutritional value of wildrice is con-
siderably better than rice having nearly twice as much protein
and an amino acid complex similar to oat groats (hulls
removed). Wildrice is often mixed with rice and served as a
blend. The pure product serves as a side dish similar to pota-
toes or rice. Continued improvement in yield/ha should make
wildrice more available in stores and restaurants not only in
the United States but also in other countries.
Wildrice Species
Wildrice was first classified by C. Linnaeus, in England in 1753.
He used the description sent to him by J. F. Gronovius, Leiden,
Holland. Gronovius described a plant sent to him by John
Clayton that he collected in Virginia in 1739. C. Linnaeus
gave the plant the binomial Zizania aquatica L. Zizania is a
small genus of aquatic grasses in the tribe Zizanieae. Rice,
Oryza sativa L., belongs to the tribe Oryzeae, which precedes
the tribe Zizanieae. Both wildrice and rice belong to the grass
family, Gramineae, and the subfamily, Poacoideae. The earlier
hierarchical classification is presented in Grays Manual of Bot-
any, eighth edition, by Fernald (1950). However, in more
recent publications Grass Systematics by Gould and Shaw in
1983, second edition, and Flowering Plants of the World by
Heywood (1993), both wildrice and rice are classed in the
family, Poaceae, subfamily Bambusoideae, and tribe, Oryzeae.
Subsequently, wildrice growing naturally in lakes and rivers
http://dx.doi.org/10.1016/B978-0-12-394437-5.00014-0

Zizania palustris varieties
palustris and interior
Zizania aquatica variety aquatica
Zizania aquatica variety brevis
Zizania texana
Figure 1 The natural distribution in shallow lakes and rivers in North America of the three species of Zizania. Reproduced from Oelke EA, Porter RA,
Grombacher AW, and Addis PB (1997) Wild rice New interest in an old crop. Cereal Foods World 42: 234247. American Association of Cereal
Chemists, Minneapolis, MN, with permission.
was classed into three different species by Linnaeus and others.
They are two annual species Z. aquatica L. and Z. palustris L. and
one perennial species, Z. texana Hitchc. The distribution of the
three species in North America is shown in Figure 1.
There are two varieties of annual Z. aquatica that grow along
the eastern Atlantic seacoast. Their seed is long and thin and
generally not harvested for food. Z. aquatica can also be found
in the Z. palustris area but to a limited extent. The two varieties
of annual Z. palustris have relatively large grains and have been
and are harvested as food by Native Americans. Today, the
most extensive natural stands occur in the states of Minnesota
and Wisconsin in the United States and in the provinces of
Manitoba and Ontario, Canada. Z. texana is a perennial and
only grows in a small stream in Texas; it has small grains and is
not harvested. Z. latifolia Turcz. is also a perennial and can be
found in Asia. There are accounts that the grain has been used
by the elite as a special food. However, today in Asia, the base
of the plant, which is infected with a fungus, is used as a
vegetable, Makomo-taki. A brief description of each species
is given in Table 1.
Historical Use of Species
Zizania palustris L.
The large seed from this species, ranging from
8 to 16 mm
long and
1.22.5 mm in diameter, has been harvested from
lakes and rivers in the Great Lakes region by the Chippewa
(Ojibwa) and Menominee Native American tribes for
THE CEREAL GRAINS | Wildrice, Zizania: Overview 131
hundreds of years. After harvest, it was processed and stored
and used as a carbohydrate source for the long, cold winters.
Many battles were fought for control of the prized wildrice
stands. In most cases, the Chippewa ended up controlling the
wildrice stands. The Ojibwa name for wildrice is Manoomin
meaning good berry or good seed, but the exact meaning is not
agreed upon. The early explorers, both English and French,
who often traded with the Ojibwa, gave various names to this
plant; some were Indian rice, wild rice, Canadian rice, squaw
rice, water oats, march oats, and water rice. The French
explorers called the plant folle avoine, fools oats, or wild
oats. The name that was most commonly used was wild rice
and is used in the trade today. To avoid confusion with the
wild species of Oryza genus, writers often use the hyphenated,
wild-rice, or the single word, wildrice. In this article, we are
using the single word wildrice. Today, stands of wildrice on
reservations in Minnesota and Wisconsin are controlled by
Native Americans, but in lakes and rivers outside of the
reservations, the stands are controlled by the Departments of
Natural Resources of each state. In Minnesota and Wisconsin, a
license is required by everyone to harvest from lakes and rivers
that are not on Native American reservations. Today, natural
stands in Minnesota are harvested, by law, in the traditional
method using a flat bottom boat or canoe. The canoe is pushed
through the stand by one individual using a long pole, while
the other individual uses two flails (knockers) to knock the
grain into the canoe. Since the grain ripens at different times on
the same panicle and easily falls (shatters) from the plant when
132 THE CEREAL GRAINS | Wildrice, Zizania: Overview

Table 1 Description of Zizania species

Species name Description

Zizania aquatica var. aquatica L. southern
wildrice
Zizania aquatica var. brevis, Fassett,
estuarine wildrice
Zizania palustris var. palustris L. northern
wildrice
Zizania palustris var. interior (Fassett),
Dore, interior wild rice
Zizania texana A.S. Hitchc., Texas wildrice
Zizania latifolia Turcz. Manchurian water
rice
Annual, tall, large panicles, many branched florets. Grows in muddy shores of streams in southern
Ontario and Quebec, southward to Florida and Louisiana. Average seed size: 14.3 mm long and 0.9 mm
diameter
Annual, short, small panicle with few branched florets. Grows in tidal areas of St. Lawrence River estuary.
Average seed size: 6.7 mm long and 1.0 mm diameter
Annual, height ranges from short to tall, slender few-flowered panicle. Grows in water up to 120 cm deep
in southern Canada from New Brunswick to Manitoba and northern US states. Average seed size:
16.7 mm long and 1.4 mm diameter
Annual, medium tall, medium to large panicle with numerous florets. Grows in water
3060 cm deep
along rivers in southeastern Manitoba and adjoining Ontario and in the North Central States. The
germplasm of this species and variety has been used to develop cultivars for cultivation. Average seed
size: 10.7 mm long and 1.8 mm diameter
Perennial, decumbent with many long stems, panicles short, short seeds. Grows only in a localized area
in San Marcos River in Texas. Average seed size: 6 mm long and 1.2 mm diameter
Perennial, spreads by subterranean runners, tall, medium panicles, seeds short to medium in length.
Native grass of Manchuria, Korea, Japan, Burma, and northeastern India. Base of plant often becomes
infected with a fungus that is used as a vegetable delicacy. This species is commercially grown in
Japan, Korea, and China for its vegetable. Average seed size of collections from Japan: 7 mm long and
1.3 mm diameter

Figure 2 The traditional way (canoe and flail) of harvesting natural stands of wildrice. One individual (standing) propels the canoe through the stand,
while the other uses two flails (knockers) to remove the grain from the stalks. Photographs by Oelke EA, left photograph appeared in Oelke EA
(2007) Saga of the Grain A Tribute to Minnesota Cultivated Wild Rice Growers. Lakeville, MN: Hobar Publications.

mature, stands are harvested by this method every other day for
a period of 2 weeks. Enough grain falls back into the water
before and during harvest to reseed the lake or river for the next
year. Figure 2 depicts the traditional harvest method. In Can-
ada, natural stands are generally harvested by airboats fitted
with a grain catcher onto the front.
The traditional method of processing the harvested wildrice
was to lay it on the ground for drying since the grain has
40%
moisture when harvested. Later, the Native Americans used an
iron kettle over a fire to parch (roast and dry) the grain
(Figures 3 and 4). The hulls (lemma and palea) were removed
from the warm grain by placing it into a round-bottom con-
tainer or leather-lined hole and then walking in place (jigging)
in the container. The chaff was removed by tossing the grain
into the air (winnowing) and allowing the wind to blow it
away. The heavier grain was caught in the container. Today,
very little of the grain harvested from natural stands is
processed in this manner. Most of it is processed with mechan-
ical equipment as described later.
Zizania aquatica L. and Zizania texana Hitchc.
The seed from these species is either too thin or small to make
it worthwhile to harvest as a food source. Z. texana is on the
endangered species list.
Zizania latifolia Turcz.
This species is a perennial and is a native grass of Manchuria,
Korea, Japan, Burma, and northeastern India. The plant gener-
ally is not used for grain production but grown commercially
for its vegetable delicacy, especially in Japan, Korea, and China.
The base of the plant becomes infected with a fungus. The
infected base of the plant is harvested and eaten as a vegetable.
In Japan, they call the vegetable Makomo-taki. The infected
part is also dried and allowed to develop spores. The spores are

Figure 3 The traditional method of drying wildrice grain in a kettle
above a fire. The grain is stirred while drying. Photograph by Oelke EA,
appeared in Oelke EA, Grava J, Noetzel D, et al. (1982) Wild Rice
Production in Minnesota. St. Paul, MN: University of Minnesota.
Agricultural Extension Service, Bulletin 464.
then used by artists to develop lacquered, antique-looking
woodwork, such as plates and small boxes (Figure 5).
Cultivation of Zizania palustris L.
Domestication Versus Cultivation
The expansion of this species of wildrice represents one of the
largest modern efforts to domesticate a cereal grain. Peperno
describes cultivation as a routine cycle of sowing and
harvesting. Native Americans did some form of cultivation
but under natural conditions. They annually harvested natural
stands relying on self-seeding of the lakes but sometimes added
seeds to lakes with poor stands. Most crops are now truly
domesticated, which means they depend mostly on human
care for their perpetuation such as annual seeding. Wildrice is
not far removed from the early cultivation phase of relying on
self-seeding. Cultivars are being developed that shatter less
than the plants growing in the lakes and rivers. They also
produce more grain per hectare but still only 1015% of that
produced by rice.
History of Cultivation/Domestication
Rice has been under the cultivation/domestication process for
some thousands of years. Interest in cultivating this plant has
been expressed for well over 100 years by businessmen and
botanists. Early explorers collected seed for planting in Europe,
THE CEREAL GRAINS | Wildrice, Zizania: Overview 133
Figure 4 The traditional way (jigging) to remove the outer hull of the
wildrice grains. The individual walks in place to rub the grain against each
other and also the kettle. Photograph by Oelke EA, appeared in Oelke EA,
Grava J, Noetzel D, et al. (1982) Wild Rice Production in Minnesota, St. Paul:
University of Minnesota. Agricultural Extension Service, Bulletin 464.
Figure 5 The base of the stem of Zizania latifolia, which is infected with
a fungus, left. This base is harvested especially in Japan and used as a
vegetable Makomo-taki delicacy, slices in frying pan. Photograph by
Oelke EA.
but they failed. In 1828, Timothy Flint in Geography and History
wondered why no attention was paid to this plant. In 1852 and
in 1853, Joseph Bowron and Oliver Kelly, respectively, also
thought about why this plant is not cultivated. Charles
Chambliss, USDA, ARS, in the early 1900s researched wildrice
as a potential crop. In 1917, H. B. Williams and Z. Durand
started mechanically harvesting private land in Canada. The
first individuals to construct a field specifically to commercially
produce this aquatic grain were James and Gerald Godward.
They grew wildrice in 0.5 ha diked field (paddy) at Bass Lake
near Merrifield, Minnesota. During the first 3 years, the God-
wards had good crops, but disease destroyed the crop in the
134 THE CEREAL GRAINS | Wildrice, Zizania: Overview
fourth year. They continued their pioneering efforts and by
1958 had a 50 ha diked field for growing wildrice. During the
mid-1950s and early 1960s, others started growing wildrice. In
1965, Uncle Bens, Inc. started contracting for production,
which was a big impetus to produce wildrice. Today, there
are
5000 ha in Minnesota and in California. A few ha are
now also being grown in Hungary, Europe.
Genetics and Breeding
Domestication of a wild species requires that plants are
selected from wild populations that maintain their grains on
the plant even after maturity (nonshattering). The grain also
needs to mature uniformly and has more grain production in
relation to vegetative production. One of the important factors
in the domestication process was the 1963 discovery, in a
growers field, of a nonshattering plant by the grower and
two University of Minnesota scientists (Figure 6). All wildrice
species are mostly cross-pollinated with the male and female
flowers separated on the same stem. Male flower retention and
seed shattering are almost always associated, facilitating selec-
tion for nonshattering. Subsequently improved cultivars were
developed from this germplasm by individuals, companies,
and the University of Minnesota. Up until that time, fields
Figure 6 The panicle on the left is a nonshattering type, while the one
on the right is a shattering type. The nonshattering type can be
recognized by loss of male flowers soon after pollen shed on the
branched (lower) portion of the panicle. Photograph by Oelke EA,
appeared in Oelke EA, Grava J, Noetzel D, et al. (1982) Wild Rice
Production in Minnesota. St. Paul, MN: University of Minnesota.
Agricultural Extension Service, Bulletin 464.
were planted with seed collected from lakes. These fields were
harvested with picker/harvesters on tracks. The fields had to be
harvested several times during a 2-week period, due to seed
shattering, without cutting off the plants. Yields with this
method were
6890 kg ha1 of processed grain. Fields
planted to the newer nonshattering cultivars could be har-
vested once with a regular grain combine and yielded over
700 kg ha1 (Figure 7). The development of the nonshattering
cultivars resulted in a tremendous expansion of cultivated
hectares in Minnesota and later in California. Figure 8 shows
the cultivated wildrice production after processing for Minne-
sota and California since 1968.
Individual wildrice growers began their own selection pro-
grams to select better plant types in the 1960s, and some are
continuing today. The wildrice breeding program at the Univer-
sity of Minnesota began in 1972. It continues till today and is
Female flowers (grain)
Male flowers
Panicle
Stem
Leaves
Tillers
Internode
section
Parchment-like
cross-partitions
Adventitious
root
Figure 7 A drawing of a Zizania palustris plant. This type is a typical
nonshattering plant cultivated in fields. Reproduced from Oelke, EA,
Elliott WA, Kernkamp MF, and Noetzel DM (1973) Commercial
Production of Wild Rice. St. Paul, MN: University of Minnesota.
Agricultural Extension Service, Folder 284.
 THE CEREAL GRAINS | Wildrice, Zizania: Overview 135

14000

MN+CA
12000
CA

MN
10000

Annual production (x1000 kg)

Lakes

8000

6000

4000

2000

0
68

70

72

74

76

78

80

82

84

86

88

90

92

94

96

98

00

02

04

06

08

10

12
19

19

19

19

19

19

19

19

19

19

19

19

19

19

19

19

20

20

20

20

20

20

20
Figure 8 The amount of processed wildrice in 1000 kg from 1968 to 2013 from lakes and rivers in Minnesota (
30%) and Canada (bottom red line),
Minnesota cultivated fields (yellow line), California cultivated fields (purple line), and the total from cultivated fields in Minnesota plus California
(top blue line). Graph by Porter RA (2014) Data provided by the Minnesota Cultivated Wild Rice Council, California Wild Rice Board, various wildrice
industry sources and Minnesota Department of Natural Resources.

the only public wildrice breeding program in the United States. up development resulting in a short, unproductive plant. The
To date, nine cultivars have been released all with some desir- high humidity also results in severe leaf disease problems.
able characteristics needed to further the domestication of wild- Cultivars have been developed that grow well in the warm day
rice. Improved shattering resistance is the primary goal, but but cooler night temperatures of Northern California. The cul-
disease resistance is a close secondary goal. Another goal is tivation practices for wildrice are similar to rice. Fields need to
shorter plants with higher yields. Yields have nearly doubled be kept flooded until several weeks before harvest. Wildrice will
from the original Johnson nonshattering (some resistance to grow well on organic or inorganic soils if nutrients are supplied.
shattering) cultivar developed from the nonshattering plants Wildrice requires less nitrogen than rice since it can get too tall
found in 1963. A private breeding program exists today in and lodge (fall down) if overfertilized. It will grow in deeper
California, which has developed cultivars for their warmer and and cooler water than rice; thus, water depth can be used as a
longer growing season. The genetic material for that program means of weed control in wildrice. Handling wildrice seed is
basically came from Minnesota. Thus far, all of the breeding is critical in establishing fields. Wildrice seed must be stored wet
done by conventional selection and not genetic engineering. and in cold (3  C) water for 3 months to release seed dormancy.
A program at Lakehead University, Thunder Bay, Canada, In Minnesota, new fields are seeded in the fall and the fields
selected types of plants that did well in specific lakes, but that flooded either in the fall or in the spring to a depth of
30 cm.
program has been reduced. Cultivated wildrice (Zizania palustris In Minnesota, fields can be kept in production for 23 years and
var. interior and palustris L.) of North America is an annual then rotated with other crops. After the first year, the fields seed
diploid with a chromosome number of 2n 2x 30; thus, it themselves from shattered seed even when initially seeded to
has 15 chromosome pairs. Compared with rice (Oryza sativa), the newer more shatter, resistant cultivars. In the Sacramento
wildrice has three more pairs of chromosomes and twice as much Valley of California, the fields are seeded in the spring of each
DNA content as rice. This research also indicated that when total year due to loss of viability of the shattered seed in the dry soil
wildrice DNA was used as a probe in southern hybridization to between seasons. Seed has to be stored in cool wet conditions
different Poaceae genera such as rice, oat, barley, wheat, and over the winter and then seeded the next spring. In the northern
maize, there was a strong hybridization signal with rice but higher elevations in the valleys between Mount Lassen and
relatively little with oat, barley, wheat, or maize. The close rela- Mount Shasta, the production practices are similar to Minne-
tionship of wildrice to rice will benefit the mapping of genes in sota (Figures 9 and 10).
wildrice. Similar probes used for mapping genes in rice have Fungal brown spot (Bipolaris sp.) is a severe disease in
been used in wildrice and some of the same genetic markers Minnesota, but not a problem in California. The wildrice
have been found to be located in the same area in wildrice as in worm (Apamea apamiformis Guenee), which is the larval stage
rice. Locating markers for important genes such as seed shatter- of noctuid moth, is the most serious insect pest in the Upper
ing has facilitated the wildrice breeding efforts for domestication. Midwest but not a problem in California. In Minnesota
and California, fields are drained
3 weeks before harvest.
Cultivation Practices Grain combines are adapted, with larger reels and often
full tracks, for harvesting wildrice. The grain has to be har-
Wildrice is well adapted to northern latitudes with its cooler vested at
2035% moisture since seed shattering will occur
climate. In southern latitudes of the United States, such as if harvested at lower moisture content. The grain is immedi-
Arkansas, shorter day lengths and higher temperatures speed ately transported to processing plants (Figure 11).
136 THE CEREAL GRAINS | Wildrice, Zizania: Overview
Processing
Processing consists of curing, parching (roasting and drying),
hulling, cleaning, scarification, and grading. When the grain
arrives at the processing plant, it is put into piles or containers
and kept for 1 or 2 days to allow for more uniform color. Then,
the grain is put into rotating, heated drums (parchers) that will
hold
225370 kg and heated for
1 h (Figure 12). The grain
is dried to
7% moisture and to
135  C. After parching, the
hot grain is passed through a huller consisting of two rubber
rollers going at different speeds. The light chaff and other
foreign material are evacuated from the kernels, and the hulled
kernels are graded with graders and gravity tables for size
(Figure 13). Sometimes before grading, the kernels are scari-
fied by passing through a tube with rubber paddles. This
removes some of the darker, impermeable outer layer of the
kernel to allow a faster cooking time (Figure 14). The parched,
graded kernels can then be stored for future packaging
(Figure 13).
Figure 9 Aerial view of northern Minnesota wildrice fields after flooding
in the spring. The main source of water is the river in the foreground.
Photograph by Oelke EA.
Figure 10 In Minnesota, wildrice fields in production for two or more years, plant populations are reduced by airboats. V-shaped blades mounted
on the rear are dropped into the water to the soil line to remove excess plants, which leaves rows of plants for production. Excessive amounts of seeds,
even from the more shatter resistant cultivars, fall onto the soil before and during harvest often producing too many plants the following year.
Photographs by Oelke EA.
Nutritional Properties
The wildrice kernel with the hulls removed is similar to other
cereals and consists of a pericarp, aleurone layer, endosperm,
and embryo. The pericarp and embryo each represent
5% of
the kernel weight. The rest, 90%, is largely endosperm and a
smaller amount of the aleurone layer. The nutritional quality is
very good for a cereal grain. It is equal to or better than other
cereals such as wheat, barley, or rice. The protein content is
about that of hard red spring wheat
1315%, while brown
rice has
8%. The sum of lysine, threonine, and methionine
content is similar to hulled oats, which is considered one of the
better cereals for humans. The mineral content of the wildrice
kernel varies but can be similar to oats. Processed wildrice
contains no vitamin A but is an excellent source of B vitamins.
Wildrice is nutritious, since it is not refined but consumed as a
whole grain (Tables 24).
Wildrice has good antioxidant properties. There is consid-
erable interest in antioxidants, which can slow diseases such as
coronary heart disease and certain types of cancer. Hydrated
wildrice mixed with ground beef has significant antioxidant
activity.
Markets and Uses
The cultivation and domestication of wildrice provided a large,
consistent supply of wildrice compared to the inconsistency of
that from natural stands. Today,
95% of the wildrice in the
marketplace comes from cultivated fields. The wildrice market
expanded rapidly during the 1960s through the 1980s primar-
ily in the blend market. Sales increased an average of 15% per
year when Uncle Bens, Inc., introduced the first packaged
blend of wildrice, long grain rice, and herbs. Many other
blends are now available including soups, vegetable-based
side dishes, and convenience foods such as dehydrated mixes
and frozen entrees. Increased use of wildrice as a pure product
as well as in blends should continue. There are many recipes
available for preparation and use of wildrice. Numerous
 THE CEREAL GRAINS | Wildrice, Zizania: Overview 137

wildrice cookbooks are available. In Minnesota, by law, the

pure product has to be labeled cultivated if it is produced in
cultivated fields and sold in Minnesota. Wildrice is a nutritious
food, and even though it is considered a gourmet food, it is not
very expensive on a per serving basis. Production of wildrice
should increase as new, better yielding cultivars and produc-
tion practices are developed.

Exercises for Revision

Why is the name wildrice used in scientific literature instead

Figure 11 Cultivated wildrice, nonshattering cultivars, being harvested
in Minnesota with a combine with full tracks needed for the soft wet
peat or mineral soils. Photograph by Hansen DL (2012) Agricultural
Experiment Station, University of Minnesota, St. Paul.
of wild rice?
What morphological features of wildrice make it different
from rice?
Why is only one species of wildrice used for harvesting the
grain and which one is it?
In North America, why did Native Americans revere
wildrice?
Describe the cultivation and domestication of plants. Is
there a difference between the two?
Do production practices differ from other cereal grains and
in what way?
How do the yields on a hectare basis of wildrice compare to
rice and why?
How would you compare the wildrice nutritional value for
humans to other cereal grains?
Is wildrice available in large enough quantity to remove it
from the specialty cereal grain category?

Exercises for Readers to Explore the Topic Further

Wildrice seeds need to be held in wet, cool conditions

Figure 12 Rotating drums heated on the bottom to dry (parch) the
wildrice grain. The drums are covered after the grain is put in to allow
some steaming of the grain. Temperature of the grain can be checked
with a laser thermometer. Modern processing is based on the principle
of traditional methods. In California, some of the wildrice grain is
parboiled. Photograph by Oelke EA.
to release dormancy; thus, better seed handling and
storage techniques are needed to facilitate growing and plant
breeding. See Further Reading references: Cardwell VB,
Oelke EA, Elliott WA (1978) Seed dormancy mechanisms
in wild rice (Zizania aquatica). Agronomy Journal 70: 481;
Oelke EA (2007) Saga of the Grain A Tribute to Minnesota
Cultivated Wild Rice Growers. Lakeville, MN: Hobar
Publications.

Figure 13 Processing is done with modern grain handling equipment in clean processing plants. Storage is done in clean warehouses before sale or
packaging. Photographs by Oelke EA.
138 THE CEREAL GRAINS | Wildrice, Zizania: Overview

Figure 14 Processed kernels from cultivated fields that were slightly scarified for faster cooking (left). Kernels without and with hulls on the right. Hulls
are removed during processing. Hand-processed kernels, from lakes and rivers, most often are lighter in color than the ones in the left photograph.
Photographs Oelke EA.

Table 2 Composition of wildrice, medium grain brown rice, hard red spring wheat, and oats in 100 g

Nutrient Unit Wildrice Brown rice, medium grain Wheat, hard red spring Oats

Water g 7.76 12.37 12.76 12.82

Protein g 14.73 7.5 15.4 16.89
Total lipid (fat) g 1.08 2.68 1.92 6.9
Fiber, total dietary g 6.2 3.4 12.2 10.6
Carbohydrates (by difference) 74.9 76.17 8.03 66.27
Sugars g 2.5 0.41
Calcium mg 21 33 25 54
Iron mg 1.96 1.8 3.6 4.72
Magnesium mg 177 143 124 177
Phosphorus mg 433 264 332 523
Potassium mg 427 268 340 429
Sodium mg 7 4 2 2
Zinc mg 5.96 2.02 2.78 3.97
Copper mg 0.524 0.277 0.41 0.626
Manganese mg 1.329 3.74 4.055 4.916

Data from US Department of Agriculture, Agricultural Research Service (2013) USDA National Nutrient Database for Standard Reference, Release 26.

To facilitate the selection of better cultivars, better and
new genetic markers are needed. See Further Reading
references: Hass BL, Pires JC, Porter R, Phillips RL (2003)
Comparative genetics at the gene and chromosome levels
between Oryza sativa and wildrice Zizania palustris.
Theoretical and Applied Genetics 107: 773782; Kennard
WC, Phillips RL, Porter RL (2002). Genetic dissection of
shattering, agronomic, and color traits in American wildrice
(Zizania palustris var. interior L.) with a comparative map.
Theoretical and Applied Genetics 105: 10751086; Kennard
WC, Phillips RL, Porter RA, Grombacher AW (2000) A
comparative map of wild rice (Zizania palustris L.
2n 2X 30). Theoretical and Applied Genetics 101:
677684.
Processing of wildrice is still a batch system that slows
processing; thus, there is a need to develop storage
techniques for the high moisture grain before processing.
See Further Reading references: Marcum DB (2007) Wild
Rice Production in California. Davis, CA: University of Cal-
ifornia. Extension Publication, No.21622; Oelke EA,
Boedicker JJ (2000) Wild rice: Processing and utilization.
In: Kulp K, Ponte JG (eds.) Handbook of Cereal Science and
Technology, pp. 275306. New York: Marcel Dekker; Oelke
EA (2007) Saga of the Grain A Tribute to Minnesota
Cultivated Wild Rice Growers. Lakeville, MN: Hobar
Publications.
Additional research needs to be conducted to verify the low
presence or absence of gluten in processed wildrice.
 THE CEREAL GRAINS | Wildrice, Zizania: Overview 139

Table 3 Amino acids in wildrice, oats, medium grain brown rice, Archibold OW (1994) Wild Rice in Saskatchewan Agricultural Development in
and hard red spring wheat in 100 g Harmony with Nature. Saskatoon: Saskatchewan Agriculture and Food.
Cardwell VB, Oelke EA, and Elliott WA (1978) Seed dormancy mechanisms in wild rice
Amino acid Unit Wildrice Oats Brown rice Wheat (Zizania aquatica). Agronomy Journal 70: 481.
Chang TT (1989) Domestication and spread of the cultivated rices. In: Hillman GC
Tryptophan g 0.179 0.234 0.096 0.195 and Harris DR (eds.) Foraging and Farming: The Evolution of Plant Exploitation,
Threonine g 0.469 0.575 0.275 0.433 pp. 408417. UK: Routledge.
Fassett NC (1924) A study of the genus Zizania. Rhodora 26: 153.
Isoleucine g 0.618 0.694 0.318 0.541
Hass BL, Pires JC, Porter R, and Phillips RL (2003) Comparative genetics at the gene
Leucine g 1.018 1.284 0.620 1.038 and chromosome levels between Oryza sativa and wildrice Zizania palustris.
Lysine g 0.629 0.701 0.286 0.404 Theoretical and Applied Genetics 107: 773782.
Methionine g 0.438 0.312 0.169 0.230 Jenks AE (1900) The wild rice gatherers of the Upper Lakes: A study in American
Cystine g 0.174 0.408 0.091 0.404 primitive economics. In: 19th Annual Report of the Bureau of American Ethnology,
Phenylalanine g 0.721 0.895 0.387 0.724 189798, Part 2, pp. 10131137. Washington, DC: US Department of Interior.
Tyrosine g 0.622 0.573 0.281 0.441 Kennard WC, Phillips RL, Porter RA, and Grombacher AW (2000) A comparative map of
Valine g 0.858 0.937 0.440 0.679 wild rice (Zizania palustris L. 2n 2X 30). Theoretical and Applied Genetics
Arginine g 1.136 1.192 0.569 0.702 101: 677684.
Kennard WC, Phillips RL, and Porter RL (2002) Genetic dissection of shattering,
Histidine g 0.384 0.405 0.190 0.330
agronomic, and color traits in American wildrice (Zizania palustris var. interior L.)
Alanine g 0.825 0.881 0.437 0.555 with a comparative map. Theoretical and Applied Genetics 105: 10751086.
Aspartic acid g 1.419 1.448 0.702 0.808 Kosbau H (2005) The Taming of Wild Rice. Grand Rapids, MN: Treasure Bay Printing.
Glutamic acid g 2.565 3.712 1.528 4.946 Marcum DB (2007) Wild Rice Production in California. Extension Publication, Davis,
Glycine g 0.672 0.841 0.369 0.621 CA: University of California, No.21622.
Proline g 0.519 0.934 0.352 1.680 Nelson RN and Dahl RP (1985) Wild Rice Market Shows Vigorous Growth. St. Paul,
Serine g 0.778 0.750 0.388 0.663 MN: University of Minnesota, Minnesota Agriculture Economics, No.649.
Oelke EA (2007) Saga of the Grain A Tribute to Minnesota Cultivated Wild Rice
Data from US Department of Agriculture, Agricultural Research Service (2013) Growers. Lakeville, MN: Hobar Publications.
National Nutrient Database for Standard Reference, Release 26. Oelke EA and Boedicker JJ (2000) Wild rice: Processing and utilization. In: Kulp K and
Ponte JG (eds.) Handbook of Cereal Science and Technology, pp. 275306.
New York: Marcel Dekker.
Table 4 The amounts of nutrients based on 1/2 cup serving of Oelke EA, Elliott WA, Kernkamp MF, and Noetzel DM (1973) Commercial Production of
cooked product Wild Rice. St. Paul, MN: University of Minnesota, Agricultural Extension, Service
Extension Folder 284.
Nutrients (based on 1/2 Enriched white Brown rice Oelke EA, Grava J, Noetzel D, et al. (1982) Wild Rice Production in Minnesota.
cup serving (cooked)) Wildrice rice (long grain) (long grain) Agricultural Extension Service, St. Paul, MN: University of Minnesota, Extension
Bulletin 464.
Calories 83 103 108 Oelke EA, Porter RA, Grombacher AW, and Addis PB (1997) Wild rice New interest in
Protein (g) 3.27 2.13 2.52 an old crop. Cereal Foods World 42: 234247, American Association of Cereal
Chemists, Minneapolis, MN.
Fat (g) 0.28 0.22 0.88
Piperno DR (2012) Archaeobotanical information on early cultivation and plant
Dietary fiber (g) 1.5 0.3 1.8
domestication involving microplant (phytolyth and starch grain) rements.
Total carbohydrates (g) 17.5 22.25 22.39 In: Gepts P, Famula TR, Bettinger RI, Brush SB, Damania AB, McGuire PE, and
Minerals Qualset CO (eds.) Biodiversity in Agriculture: Domestication, Evolution, and
Phosphorus (mg) 67 34 81 Sustainability, p. 203. Cambridge: Cambridge University Press.
Iron (mg) 0.49 0.95 0.41 Steeves TA (1952) Wild rice Indian food and a modern delicacy. Economic Botany
Potassium (mg) 83 28 42 26: 107142.
Zinc (mg) 1.1 0.39 0.61 Vennum T (1988) Wild Rice and the Ojibway People. St. Paul, MN: Minnesota Historical
Vitamins Society.
Wu K, Zhang W, Addis PB, Epley RJ, Salih A, and Lehrfeld J (1994) Antioxidant
Riboflavin (mg) 0.07 0.01 0.02
properties of wild rice. Journal of Agricultural and Food Chemistry 42: 34.
Niacin (mg) 1.05 1.16 1.49
Vitamin E (mg) 0.2 0.03 0.03

Reproduced with permission from the Minnesota Cultivated Wild Rice Council, St. Paul.
Data from the U. S. Department of Agriculture, Agricultural Research Service (2013)
USDA National Database for Standard References, Release 26.
See also: Breeding of Grains: Rice: Breeding; Genetics of
Grains: Rice: Genetics; Non-wheat Foods: Rice: Chinese Food
Uses; The Basics: Taxonomic Classification of Grain Species; The
Cereal Grains: An Overview of the Family of Cereal Grains Prominent
in World Agriculture; Oats: Overview.
Further Reading
Aiken SG, Lee PF, Punter D, and Stewart JM (1988) Wild Rice in Canada. Winnipeg:
Agriculture Canada.
Relevant Websites
http://www.ars.usda.gov/ba/bhnrc/ndl This website is an excellent source for nutrient
content of many grains and foods including wildrice.
http://www.cawildrice.com/ This is the website of the California Wild Research and
Advisory Board of the California cultivated wild rice growers. Information on
California wildrice cultivation, food preparation and recipes can be found on this
website.
http://www.mnwildrice.org This is the website of the Minnesota Cultivated Wild
Rice Council whose members consist of Minnesota cultivated wildrice
growers. Information on wildrice history, Minnesota cultivation, research,
processing, nutrition, gluten content, cooking, and recipes can be found on this
website.
http://www.wildrice.org This website is the one for the International Wild Rice
Association (IWRA). IWRA consists of wildrice growers, processors and marketers.
Members include individuals from the US and Canada. Information on wildrice
history and links to other wildrice websites can be found on this website.
 African Rice (Oryza glaberrima): A Brief History and Its Growing Importance
in Current Rice Breeding Efforts
JT Manful and S Graham-Acquaah, Africa Rice Center, Cotonou, Republic of Benin
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The two cultivated species of rice are Oryza sativa (Asian
rice) and Oryza glaberrima (African rice).
O. glaberrima have tolerance to several biotic and abiotic
stresses and are particularly suited for low-input subsistence
farming as practiced in many African communities.
African rice has yield potential comparable to O. sativa as
evidenced by yield-related morphoagronomic traits. Their
tendency to easily shatter is a responsible factor for the
lower yields observed.
NERICA (New Rice for Africa) a product of interspecific
breeding between O. glaberrima and O. sativa developed by
the Africa Rice Center combines the stress tolerance of
African rice with the high yield potential of Asian rice.
The recent release of O. glaberrima genome and the devel-
opment of African rice-specific single-nucleotide polymor-
phisms will enhance the identification and exploitation of
adaptive traits in African rice in rice improvement
programs.
Very little work has been done on the physicochemical
properties of O. glaberrima. Ongoing studies by the Africa
Rice Center show that a high level of genetic diversity exists
in the accessions of O. glaberrima from the West Africa sub-
region as evidenced by variations in both apparent amylose
content and pasting properties.
Learning Objectives
At the end of this article, it is expected that readers would
have knowledge of the origin, domestication history, and
diversity in African rice;
understand the traits that set O. glaberrima apart from O.
sativa;
know how O. glaberrima is used in classical breeding;
have an appreciation of ongoing research work on the
physicochemical properties of African rice.
Introduction
Rice is a major staple in the world and belongs to the genus
Oryza. Out of over 20 species in the genus Oryza, only two
species are cultivated: Oryza glaberrima (African rice) and Oryza
sativa (Asian rice). O. glaberrima is native to sub-Saharan Africa
(SSA) thought to have been domesticated from the wild ances-
tor Oryza barthii (formerly known as Oryza breviligulata).
O. glaberrima is presently being replaced everywhere in
Africa by O. sativa and is rapidly diminishing in importance.
Recent advances now allow for gene transfer between the two
140 Encyclopedia of Food Grains, Second Edition
species, thus creating crosses that are better adapted, and
higher yielding under adverse conditions, than either parent.
This article discusses the history of glaberrima, its genetic and
morphoagronomic diversity, its use in conventional breeding,
and some of its physicochemical characteristics.
Origin and History of O. glaberrima
O. glaberrima is unique to Africa. It is believed to have been
domesticated from the wild ancestor O. barthii over 3000 years
ago. The many indigenous words for rice in several West Afri-
can language groups predate the Portuguese-derived words
associated with Asian rice and offer linguistic evidence to the
fact that rice was cultivated prior to the arrival of Europeans in
Africa. Two distinct theories have been advanced as to the exact
origin and domestication of the African species. The first theory
propounded by Porteres suggests that O. glaberrima was first
domesticated in the Inland Delta of the Niger River in present-
day Mali and then subsequently spread to other centers of
diversification, some being the coast of Gambia, the Casa-
mance region of Senegal, and Guinea Bissau along the Niger
River and then in the Guinea forest between Sierra Leone and
the western Ivory Coast. Others claim that ancient hunting and
gathering human populations living in different localities
within the vast forest and savanna areas where O. barthii grew
may have selected for O. glaberrima. Recent studies into the
population structure and genomics of O. glaberrima accessions
tend to support a single domestication origin of African rice in
areas of the upper Niger and Senegal Rivers.
African rice, as has been asserted, was introduced into the
new world via the trans-Atlantic slave trade. Botanical evidence
and historical records suggest that African rice was grown in El
Salvador, French Guiana, Brazil, Jamaica, South Carolina, and
Suriname and is to this day grown, eaten, and exported by the
Maroons. Although it has diminished in terms of cultivated
area as a result of the introduction of O. sativa, O. glaberrima
remains an important crop for certain communities in West
Africa who still grow them for use in rituals and as food due to
its unique taste and texture. Their hardiness in the field also
makes them ideal for subsistence farming.
What Makes O. glaberrima Different?
Differentiating between the two cultivated species of rice in the
field can be cumbersome for the untrained as they differ mar-
ginally in their morphology. In general, O. glaberrima has short
round ligules, while the panicles are erect and simply branched
with small spikelets. The grains are small and pear-shaped with
a reddish-brown to purple bran color and olive to black seed
coats. As a result of this bran color, O. glaberrima are often
http://dx.doi.org/10.1016/B978-0-12-394437-5.00016-4
 THE CEREAL GRAINS | African Rice (Oryza glaberrima): History and Its Growing Importance in Rice Breeding Efforts
Plate 1 Panicles of O. sativa (a) and (b) and O. glaberrima (c) and (d).
referred to as red rice. Compared with Asian varieties, African
rice usually gives lower grain yields, readily shatters, and is
difficult to mill because of the brittle nature of the grains
(Plate 1).
These notwithstanding, O. glaberrima possess distinct char-
acteristics, perhaps due to the fact that they have survived with
little interference from humans, which make them ideal for the
low-input subsistence farming still practiced in many African
communities. The plants have wider leaves that shade out
weeds and are more tolerant than O. sativa to biotic and abiotic
stresses such as diseases, pests, fluctuations in water depth, iron
toxicity, infertile soils, severe climatic conditions, and neglect.
Diversity in African Rice
O. glaberrima is largely being replaced by O. sativa because of
the higher yields obtained from the latter. In spite of this, it is
still being grown in many areas because of their preference for
the preparation of certain traditional foods and also for their
tolerance to biotic and abiotic stresses. This stress tolerance
prompted efforts at its conservation and use as a gene pool
for improving Asian rice varieties. The Africa Rice Center
(www.africarice.org) is leading efforts at conservation and
exploitation of O. glaberrima in breeding. The gene bank of
the center contains about 3000 accessions of O. glaberrima that
are being systematically screened.
O. glaberrima has long been perceived to possess less genetic
diversity in comparison with O. sativa. In fact, African rice is
reported to have only one-sixth of the nucleotide diversity
found in Asian rice. Indica rice (O. sativa ssp. indica) alone
has higher diversity (ysil 0.00280.0029) than either African
rice (ysil 0.0007) or its wild ancestor O. barthii (ysil 0.0024)
(Wattersons estimator of y per base pair calculated based on
141
silent sites). African rice is reported to have more than 70% less
diversity compared to its wild progenitor resulting from a
severe genetic bottleneck during its domestication from the
initial population of O. barthii.
An earlier characterization of 198 accessions using SSR
(simple sequence repeat) markers showed a range of 227
alleles per locus with an average of 9.4 alleles per locus.
Based on this characterization, O. glaberrima were grouped
into five genetic groups with two of these segregating with O.
sativa indicating intermediaries and thus gene flow between the
two species.
A morphoagronomic and molecular characterization of
African rice varieties collected from the inland delta of the
Niger River (Mopti and Tombouctou), Sikasso, and Gao
Region in Mali believed to be centers of domestication for O.
glaberrima revealed a higher average heterozygosity (0.18) in O.
glaberrima than previously reported. The population collected
from Mopti and Gao Region had higher diversity than other
populations. Although there was no clear clustering of individ-
uals based on location or growing condition because of the
probable higher gene flow that occurs between different grow-
ing conditions within a location, a higher differentiation was
observed between populations from different locations than
from different ecologies, while within locations, similarities
were observed between populations from similar growing con-
ditions. The accessions RAM116, RAM100, and RAM122 out-
performed the best check varieties under drought conditions.
Recently, single-nucleotide polymorphisms (SNPs) specific
to African rice have been developed and used to genotype some
279 O. glaberrima accessions, selected based on their impact in
current breeding programs and their geographic distribution,
and 101 O. barthii accessions, selected based on their
geographic origins. A structuring of accessions into three popu-
lations of O. barthii associated with geography and two
142 THE CEREAL GRAINS | African Rice (Oryza glaberrima): History and Its Growing Importance in Rice Breeding Efforts

populations of O. glaberrima that are yet to be linked with any
phenotyped traits was reported. Also, an apparent misclassifi-
cation in both O. glaberrima and O. barthii was identified in
addition to a new form of O. sativa within the African rice
varieties. The complete genomic sequence of O. glaberrima
has been deciphered through immense global research effort.
The release of O. glaberrima genome and the development of
African rice-specific SNPs bring in their wake hitherto non-
existing opportunity for the identification and exploitation of
adaptive traits in African rice in rice improvement programs.
Little variation has been reported in the morphoagronomic
traits of O. glaberrima compared with O. sativa. With respect to
their vegetative growth, more than 90% of O. glaberrima
screened by the Africa Rice Center exhibited high seedling
vigor. A very rapid vegetative growth culminating in large
number of tillers and droopy lower leaves was observed.
These vegetative growth characteristics (vigorous initial
growth) typified by their high tillering ability, large leaf area,
and droopy leaves provided rapid ground cover, thus suppres-
sing weeds and making them weed-competitive (Plate 2).
O. glaberrima also vary greatly in their growth durations
with as much as 89 days between some varieties. Whereas
traditional O. sativa and improved varieties have maturity
days ranging between 120140 days and 80120 days, respec-
tively, some O. glaberrima can mature within 75 days. Such
early-maturing varieties ensure that they escape drought in
drier areas and late-season fungal disease in wet ecologies.
Rice plant height and stem strength are related to lodging.
Generally taller varieties with weaker stems are more prone to
lodging especially in high soil fertility conditions. Most O.
glaberrima varieties are deemed intermediate in stature
(80100 cm) but possess fragile culms that render them sus-
ceptible to lodging during their vegetative stage. A number of
O. glaberrima have, however, been observed to have thick leaf
sheaths making them resistant to lodging. Such varieties may
be useful in breeding.
The greatest limitation to the cultivation of O. glaberrima is
their lower yields. Yield-related characteristics have been
studied, and results point to the fact that O. glaberrima have
the same yield potential as O. sativa. Yield-related morpho-
agronomic traits such as average panicle number in O. glaber-
rima are comparable to O. sativa. Shattering could also be a
contributory factor to the inferior yields as spikelet number in
O. glaberrima prior to shattering is no less than in O. sativa. O.
glaberrima possess higher dry matter production and greater
leaf area than O. sativa at least until heading. They have lower
photosynthetic rate per leaf area but a higher rate against the
same leaf nitrogen content in a low content range. Their
response to dry matter, leaf area, and leaf photosynthesis to
increases in nitrogen inputs is higher than in O. sativa.

AG: Vigor of selected O. glaberrima and O. sativa after 14 days of seeding dry season

AG: Vigor of selected O. glaberrima and O. sativa after 21 days of seeding wet season
Plate 2 Seedling vigor in O. sativa and O. glaberrima. Photo courtesy of Venuprasad Ramaiah; Africa Rice Center.
 THE CEREAL GRAINS | African Rice (Oryza glaberrima): History and Its Growing Importance in Rice Breeding Efforts
Additionally, water use efficiency on dry matter accumulation
and gas exchange bases is lower compared with that of O.
sativa, while both leaf senescence during maturity and grain
filling are faster in O. glaberrima than O. sativa.
The greatest attribute of O. glaberrima is their resilience.
Accessions of O. glaberrima have been found to be highly resis-
tant to pests and diseases such as African rice gall midge, the rice
yellow mottle virus, and blast. O. glaberrima can limit water loss
through stomata closure and leaf curling. This trait in addition
to the well-developed root system makes them well adapted to
drought conditions. The ability for strong leaf elongation under-
water that leads to enhanced expansion of leaf area and photo-
synthesis, requirements for escaping complete submergence
underwater over a prolonged period (up to 37 days), has also
been observed in O. glaberrima, making them a promising res-
ervoir of genes for selecting and breeding rice genotypes for
different flood-prone environments in Africa. Tolerance to iron
toxicity and saline environments has also been documented in
O. glaberrima. Although some O. sativa varieties are better at
tolerating some specific stresses, a unique trait in O. glaberrima
is that some can possess multiple tolerances to various stresses.
CG 14 is one such O. glaberrima variety that has weed
competiveness and iron toxicity and drought tolerance. These
make them ideally suited for cultivation in SSA where rice
farming is predominantly carried out by resource-poor farmers.
O. glaberrima in Classical Plant Breeding
At the Africa Rice Center (AfricaRice), attempts have been made
to develop new interspecific rice varieties from crosses between
O. sativa and O. glaberrima. Initially, this proved difficult as the
progenies were sterile. This problem was, however, successfully
overcome, and the first large-scale hybridization of the two
cultivated species of rice occurred. The resulting varieties were
collectively named NERICA (New Rice for Africa). These varie-
ties, developed for both upland and lowland ecologies, combine
the high yield of O. sativa with the stress tolerance and adapt-
ability to local growing conditions of O. glaberrima. The NERICA
development process began with an initial screening of 1721
accessions comprising 1130 O. glaberrima and 591 O. sativa
based on their morphoagronomic performance. O. glaberrima
(CG 14) and three O. sativa ssp. japonica (WAB56-104, WAB56-
50, and WAB181-18) are the parental lines for the first set of
NERICAs (NERICA 118). These varieties were crossed to obtain
interspecific lines that were backcrossed and the progenies sub-
jected to pedigree selection to obtain the first NERICAs (NERICA
118) for upland ecology. Further research and improved meth-
odology led to the release of 60 varieties (NERICA L1L60) for
lowlands. The NERICA-S lines have also been bred for the
Sahelian zones of West Africa. These NERICA varieties devel-
oped and disseminated have affected farmers livelihoods and
contributed to poverty reduction in SSA.
Physicochemical Properties of O. glaberrima
The physicochemical properties of rice such as milling recov-
eries, physical appearance (grain dimensions and chalkiness),
aroma, cooking and pasting properties are important as they
143
determine the market value and reputation of rice varieties.
Hitherto, studies on the physicochemical properties of O. gla-
berrima have not received as much attention as work on their
agronomic traits. The few studies conducted suggest that gen-
erally, O. glaberrima tend to break easily during milling; thus,
head rice recovery is lower compared with O. sativa. The red
pericarps of African rice make them less attractive when
polished. O. glaberrima is preferred by certain groups of con-
sumers, particularly in regions where they are still cultivated,
because of their distinct taste and flavor. Some aroma com-
pounds such as 4-ethylphenol (musty) and (E, E)-2,4-
heptadienal (fruity) have been found to be unique to
O. glaberrima. On cooking, O. glaberrima have firmer texture,
perhaps due to their higher apparent amylose content (AAC)
and unique pasting properties (low peak, low breakdown, and
higher setback viscosities) compared with O. sativa. Asian rice
however has greater diversity in AAC than African rice. While
waxy varieties exist among Asian rice, waxy varieties are yet to
be encountered among O. glaberrima.
With the broad objective of contributing to better under-
standing the physicochemical properties of O. glaberrima,
AfricaRice and its partners characterized over 1000 O. glaber-
rima accessions from its gene bank based on data of their Rapid
Visco Analyser (RVA) pasting characteristics and AAC. Based
on the AAC and RVA pasting parameters of the rice flours, the
accessions were hierarchically clustered into five groups C I,
C II, C III, C IV, and C V (Figure 1). The distribution pattern
indicated that the maximum number of accessions (282) was
included in C I, followed by C IV (242), C V (231), C III (161),
and C II (104), with all five clusters showing a large diversity of
AAC and pasting properties.
Cluster I had 282 accessions and constituted 27.6% of the
entire population. These accessions were further grouped into
five subclusters SC IA, SC IB, SC IC, SC ID, and SC IE with a
population of 44 (15.6%), 84 (29.8%), 107 (37.9), 26 (9.2%),
and 21 (7.4%), respectively. All the subclusters under cluster I
had similar ranges of AAC all of them fell under high-amylose
rice category. However, their pasting characteristics differed
greatly as shown in Table 1. Wide variations were observed for
peak and trough viscosities in all the five subclusters. With
respect to breakdown viscosity, while SC IB, SC IC, SC ID, and
SC IE were close, that of SC IA was relatively very distinct. SC IA,
SC IC, and SC IE were likewise similar with respect to final and
setback viscosities, but SC IB and SC ID showed diversity.
Cluster II with 104 accessions composed the lowest propor-
tion (10.2%) of the entire population. The broadest ranges of
peak, trough, setback, and final viscosities were observed in this
cluster. However, it had the second broadest AAC range and the
narrowest pasting temperature range. The range of AAC for
accessions in C II conferred them to intermediate and high-
amylose types of rice. Three subclusters SC IIA, SC IIB, and SC
IIC were formed under cluster II with populations of 26 (25%),
54 (52%), and 24 (23%) accessions, respectively. SC IIA and SC
IIB had similar ranges of AAC, with SC IIC having the broadest
range (Table 1). SC IIC also had the broadest range of peak,
final, setback, and trough viscosities, while SC IIB had the
narrowest range as regards peak, final, and trough viscosities.
With 161 accessions, cluster III constituted 15.8% of the
entire population of the O. glaberrima analyzed (Table 1) and
had five subclusters, namely, SC IIIA, SC IIIB, SC IIIC, SC IIID,
144 THE CEREAL GRAINS | African Rice (Oryza glaberrima): History and Its Growing Importance in Rice Breeding Efforts

O. glaberrima
accessions

C I C II C III C VI CV
(N = 282) (N = 104) (N = 161) (N = 242) (N = 231

SC IIIA SC IIIB SC IIIC SC VA SC VB SC VC

#SC IA SC IB SC IC (N = 37) (N = 39) (N = 43) (N -59) (N -35) (N = 76)
(N = 44) (N = 84) (N = 107)
SC IIID SC IIIE SC VD SC VE
SC ID SC IE
(N = 18) (N = 24) (N = 21) (N = 40)
(N = 26) (N = 21)
SC IVA SC VIB SC VIC
SC IIA SC IIB SC IIC (N = 80) (N = 94) (N = 42)
(N = 26) (N = 54) (N = 24)
SC IVD SC IVE
(N = 13) (N = 13)

Figure 1 Graphical representation of distribution of 1020 African rice accessions into 5 clusters and 23 subclusters.

and SC IIIE, making up 23%, 24.2%, 26.7%, 11.2%, and
14.9% of the accessions, respectively. Cluster III had the nar-
rowest ranges of peak, trough, setback, and final viscosities.
The accessions in this cluster belonged to the category of
intermediate- to high-amylose rice based on their AAC range.
SC IIIA had the broadest apparent amylose range, SC IIID had
the narrowest range, and that of SC IIIB and SC IIIE was
similar. SC IIIA also had the broadest ranges with respect to
peak, trough, breakdown, and final viscosities (Table 1).
Despite having the narrowest final viscosity range, SC IIIC
had the second broadest ranges of apparent amylose, peak
viscosity, and breakdown viscosity (Figure 2).
Accessions in cluster IV had the broadest range of apparent
amylose, the second broadest peak viscosity range, and the
fourth broadest ranges of both final and setback viscosities
but the narrowest range of breakdown viscosity. Low-,
intermediate-, and high-amylose rice types were all included
in the accessions within this cluster. Cluster IV formed 23.7%
of the entire population analyzed (242 accessions) and con-
tained five subclusters, namely, SC IVA, SC IVB, SC IVC, SC
IVD, and SC IVE, which constituted 33.1%, 38.8%, 17.4%,
5.4%, and 5.4% of the accessions, respectively (Table 1).
Whereas SC IVA, SC IVB, and SC IVC shared some similarities
in AAC and peak time, very diverse peak, trough, breakdown,
final, and setback viscosities were observed in all the five sub-
clusters except in SC IVE, which, though distinct with respect to
breakdown viscosity, showed very close similarity to SC IVB
with regard to peak, breakdown, final, and setback viscosities.
The 231 accessions in cluster were intermediate to high-
amylose rice types and formed 22.6% of the total number of
accessions analyzed. This cluster also contained five sub-
clusters, namely, SC VA, SC VB, SC VC, SC VD, and SC VE,
which constituted 25.5%, 15.2%, 32.9%, 9.1%, and 17.3% of
accessions, respectively (Table 1). Very similar AAC ranges
were observed in the subclusters except for SC VE. The acces-
sions in cluster V also had the broadest range of final viscosity,
second broadest range of setback and breakdown viscosities,
and the third broadest AAC range.
Conclusions
The tremendous amount of work on O. glaberrima at AfricaRice
notwithstanding the full potential of the species is yet to be
exploited. There remains the need to consider intraspecific
breeding within O. glaberrima and ways in which the multiple
resistance traits of O. glaberrima could be transferred into new
varieties.
The studies of the physicochemical properties supported
the idea that there is a high level of genetic diversity in the O.
glaberrima accessions as evidenced by the variation in both
AAC and pasting properties in and within the clusters and
subclusters. This suggests that there may be a valuable resource
of potentially very important gene pool that may be of benefit
to interested rice breeding programs.
Additionally, a detailed molecular analysis of a representa-
tive sample of these accessions from the clusters such as those
with a high tendency to retrograde (high setback and poten-
tially slower rate of digestion due to retrogradation) or form
viscous paste/gel after cooking and cooling (high final viscos-
ities) will ensure the use of the diversity in O. glaberrima in the
future development of acceptable commercial cultivated rice
varieties that have the desired amylose content and pasting
characteristics to meet consumer preferences in Africa and
elsewhere.
Exercises for Revision
Outline the theories of domestication of African rice.
How do the two cultivated species of rice differ from each
other?
Table 1 Distribution of 1020 O. glaberrima rice accessions obtained through cluster analysis using SPSS

Range

Cluster distribution PTemp

(N) AAC (%) PV (RVU) TV (RVU) BDV (RVU) FV (RVU) SBV (RVU) PT (s) ( C)

I C I (282) 24.729.4 67.3123.8 66.7124.2 0.4to 6.6 149.6296.8 71.7173.0 5.97.0 89.694.5
SC IA (44) 25.329.4 76.8115.6 73.3113.4 1.06.6 160.1225.4 71.7124.5 5.96.5 90.793.6
SC IB (84) 24.827.9 67.387.4 66.787.3 0.4 to 2.2 149.6188.3 75.0104.3 6.17.0 91.994.0
SC IC (107) 24.728.5 76.3101.6 76.6101.8 0.4 to 0.8 170.4234.9 75.8139.5 6.37.0 91.994.5
SC ID (26) 26.029.0 92.9123.8 93.0124.2 0.3 to 2.2 217.7296.8 118.0173.0 6.37.0 91.593.6
SC IE (21) 25.427.7 89.4107.9 89.7105.8 0.2 to 2.2 176.3217.1 86.8116.8 6.47.0 89.692.8
II C II (104) 20.629.6 152.2230.7 113.0183.5 6.049.5 246.5421.5 94.9224.8 5.75.9 86.491.9
SC IIA (26) 23.029.2 137.8182.7 125.3160.9 6.032.4 308.1394.8 54.8224.8 5.76.2 88.391.9
SC IIB (54) 23.929.6 135.5165.8 115.9143.0 6.934.2 246.5327.3 95.3170.7 5.75.9 87.291.0
SC IIC (24) 20.628.5 152.2230.7 113.0183.5 21.949.5 247.0421.5 94.9190.8 5.75.9 86.489.2
III C III (161) 21.726.8 80.9125.5 80.4122.2 0.2 to 159.9258.8 68.3149.8 5.97.0 89.694.0
12.8
SC IIIA (37) 21.726.8 89.2125.5 88.2122.3 0.2 to 228.3258.8 120.8149.8 5.97.0 90.793.6
12.8
SC IIIB (39) 23.226.6 89.8110.8 89.8103.1 0.2 to 9.2 212.1232.8 107.8132.2 5.97.0 90.793.6
SC IIIC (43) 22.626.5 80.9110.0 80.4104.1 0.2 to 194.3211.5 92.1119.1 5.97.0 89.994.0
11.8
SC IIID (18) 24.626.4 108.2121.3 100.7115.5 2.410.1 213.6236.1 103.3117.2 5.96.2 89.692.0
SC IIIE (24) 23.426.4 82.9107.9 81.0105.0 0.16.5 159.9196.6 68.3101.2 6.07.0 90.392.4
IV C IV (242) 15.128.6 36.493.1 36.593.0 0.5 to 1.2 70.8205.8 33.0122.0 6.37.0 91.595.6
SC IVA (80) 22.427.8 53.673.4 54.073.8 0.5 to 0.0 119.8158.4 57.989.8 7.07.0 92.895.2
SC IVB (94) 15.128.6 63.683.9 63.784.0 0.4 to 0.2 135.8180.4 62.3105.0 6.87.0 91.594.7
SC IVC (42) 23.027.6 36.463.3 36.563.4 0.4 to 0.1 70.8118.7 33.067.8 7.07.0 92.495.6
SC IVD (13) 21.327.4 74.593.1 74.393.0 0.2 to 0.4 171.0205.8 81.2122.0 6.67.0 92.094.4
SC IVE (13) 21.725.6 50.883.3 50.482.9 0.4 to 1.2 123.4176.0 60.498.0 6.36.7 91.994.0
V C V (231) 21.028.8 96.3152.3 87.1141.4 0.822.2 194.2343.5 84.8197.7 5.76.4 87.192.4
SC VA (59) 25.328.8 117.3152.3 104.9139.3 6.322.2 212.4275.3 84.8137.7 5.76.0 87.191.1
SC VB (35) 25.728.8 114.3151.1 108.5141.4 1.614.8 255.7343.5 135.2197.7 5.86.3 89.992.4
SC VC (76) 25.528.5 100.8131.6 198.3124.8 0.810.1 209.7268.1 91.3153.2 5.96.4 89.192.4
SC VD (21) 25.628.1 96.3117.2 87.1104.2 3.122.0 194.2229.4 90.5117.3 5.76.0 89.391.5
SC VE (40) 21.027.5 108.3141.5 95.9127.5 5.717.0 237.5306.7 111.4167.7 5.76.2 88.792.0

C, cluster; SC, subcluster; PV, peak viscosity; TV, trough viscosity; BDV, breakdown viscosity; FV, final viscosity; SBV, setback viscosity; PT, peak time; PTemp, peak temperature;
AAC, apparent amylose content.

TOG 14365 (glaberrima) RAM 123 (glaberrima)

TOG 12440 (glaberrima) IRGC 86741 (glaberrima)
Basmati Koshihikari
4000 Wita 4 Temperature (C)
95
3500
85
3000

75
Temperature (C)

2500
Viscosity (cP)

65
2000

55
1500

1000 45

500 35

0 25
0
36
68
100
132
164
196
228
260
292
324
356
388
420
452
484
516
548
580
612
644
676
708
740

Time (s)
Figure 2 RVA profile of selected O. glaberrima and O. sativa varieties.
146 THE CEREAL GRAINS | African Rice (Oryza glaberrima): History and Its Growing Importance in Rice Breeding Efforts
Discuss why O. glaberrima is particularly suited to low-input
subsistence farming.
Provide reasons to support the assertion that O. glaberrima
could have the same yield potential as Asian rice.
Why is O. glaberrima an important food security crop in
Africa?
Briefly describe the variations in the physicochemical prop-
erties of O. glaberrima.
Exercises for Readers to Explore the Topic Further
What are the prospects for intraspecific breeding within O.
glaberrima and how can the multiple trait resistance of O.
glaberrima be transferred into new varieties.
Does the nutritional profile of O. glaberrima differ from O.
sativa and can we derive functional foods from African rice?
See also: Barley, Rice and Maize Processing: Rice Processing:
Beyond the Farm Gate; Breeding of Grains: Rice: Breeding;
Carbohydrates: Glycemic Index, Glycemic Load, and Their Health
Benefits; Food Grains and Well-being: Functional Foods:
Overview; Genetics of Grains: Rice: Genetics; Processing of
Grains: Grain Quality Attributes for Cereals Other than Wheat; The
Cereal Grains: Cereals: Domestication of the Cereal Grains; Rice:
Overview; Wildrice, Zizania: Overview.
Further Reading
Agnoun YB, Samadori SH, Sie M, Vodouhe RS, and Ahanchede A (2012) The African
rice Oryza glaberrima Steud: Knowledge distribution and prospects. International
Journal of Biology 4(3). http://dx.doi.org/10.5539/ijb.v4n3p158.
Chang TT, Marciano AP, and Loresto GC (1977) Morpho-agronomic variousness and
economic potentials of Oryza glaberrima and wild species in the genus Oryza. Paper
presented at the Meeting on African Rice Species.
Futakuchi K, Fofana M, and Sie M (2008) Varietal differences in lodging resistance of
African rice (Oryza glaberrima Steud.). Asian Journal of Plant Sciences 7(6):
569573.
Futakuchi K and Sie M (2009) Better exploitation of African rice (Oryza glaberrima
Steud.) in varietal development for resource-poor farmers in West and Central
Africa. Agricultural Journal 4(2): 96102.
Futakuchi K, Sie M, and Saito K (2012) Yield potential and physiological and
morphological characteristics related to yield performance in Oryza glaberrima
Steud. Plant Production Science 15(3): 151163.
Gayin J, Chandi GK, Manful J, and Seetharaman K (2015) Classification of rice based
on statistical analysis of pasting properties and apparent amylose content: The case
of Oryza glaberrima accessions from Africa. Cereal Chemistry 92(1): 2228.
Jones MP, Mande S, and Aluko K (1997) Diversity and potential of Oryza glaberrima
Steud in upland breeding. Breeding Science 47: 395398.
Li ZM, Zheng XM, and Ge S (2011) Genetic diversity and domestication history of
African rice (Oryza glaberrima) as inferred from multiple gene sequences.
Theoretical and Applied Genetics 123(1): 2131.
Linares OF (2002) African rice (Oryza glaberrima): History and future potential.
Proceedings of the National Academy of Sciences of the United States of America
99(25): 1636016365. http://dx.doi.org/10.1073/pnas.252604599.
Manners G (2001) NERICA: The new rice transforming agriculture for West Africa.
Science in Africa http://www.scienceinafrica.com/old/index.php?qnerica.htm.
Ndjiondjop MN, Cisse F, Girma G, et al. (2010) Morpho-agronomic and molecular
characterisation of Oryza glaberrima germplasm from Mali. African Journal of
Biotechnology 9(44): 74097417.
Orjuela J, Sabot F, Cheron S, et al. (2014) An extensive analysis of the African rice
genetic diversity through a global genotyping. Theoretical and Applied Genetics
127: 22112223. http://dx.doi.org/10.1007/s00122-014-2374-z.
Sahrawat KL and Sika M (2002) Comparative tolerance of Oryza sativa and O.
glaberrima rice cultivars for iron toxicity in West Africa. International Rice Research
Notes 27(2): 3031.
Sakagami JI, Joho Y, and Ito O (2009) Contrasting physiological responses by cultivars
of Oryza sativa and O. glaberrima to prolonged submergence. Annals of Botany
103: 171180. http://dx.doi.org/10.1093/aob/mcn201.
Semon M, Nielsen R, Jones MP, and McCouch SR (2005) The population structure of
African cultivated rice Oryza glaberrima (Steud.) evidence for elevated levels of
linkage disequilibrium caused by admixture with O. sativa and ecological
adaptation. Genetics 169(3): 16391647.
Sie M, Sanni K, Futakuchi K, et al. (2012) Towards a rational use of African rice (Oryza
glaberrima Steud.) for breeding in Sub-Saharan Africa. Genes, Genomes and
Genomics 6(Special Issue 1): 17.
Van Andel T (2010) African rice (Oryza glaberrima Steud.): Lost crop of the enslaved
Africans discovered in Suriname. Economic Botany 64(1): 110.
Wang M, Yu Y, Habere G, et al. (2014) The genome sequence of African rice (Oryza
glaberrima) and evidence for independent domestication. Nature Genetics
46: 982988. http://dx.doi.org/10.1038/ng.3044.
Watanabe H, Futakuchi K, Jones MP, and Sobambo BA (2006) Grain protein content of
interspecific progenies derived from the cross of African rice (Oryza glaberrima
Steud.) and Asian rice (Oryza sativa L.). Plant Production Science 9(3): 287293.
Watanabe H, Futakuchi K, Teslim I, and Sobambo BA (2002) Milling characteristics and
grain quality traits of interspecific progenies of Asian rice (Oryza sativa) with African
rice (Oryza glaberrima). Japanese Journal of Tropical Agriculture 46: 4755.
 Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses
AM Stanca, University of Modena and Reggio Emilia, Reggio Emilia, Italy
A Gianinetti, F Rizza, and V Terzi, Consiglio per la ricerca in agricoltura e lanalisi delleconomia agraria, CRA-GPG, Fiorenzuola
dArda, Italy
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Barley was domesticated from its wild relative Hordeum
vulgare subsp. spontaneum in the Fertile Crescent about
10 000 years ago.
Barley is a model organism for genetic and genomic studies
in small grain cereals because it is a true diploid
(2n 2x 14).
Barley is the fourth most important cereal crop in the world.
Today, barley is used as grain for livestock feed (8090%)
and as malt for beer and other alcoholic beverages (10%),
and a small fraction is used as human food.
Specific characteristics of barley grain composition allow
barley flour to be used for the production of functional
food due to the high content of beta-glucan (48%).
Barley, in comparison with the other cereals, shows high
genetic variability for positive adaptation to environmental
stresses and growth in extreme climatic conditions.
Learning Objective
To achieve an overview of the botany, physiology, genetics,
and agronomy of barley and of the main uses of this small
grain cereal.
Barley Domestication and Spread
Barley, one of the earliest domesticated plants, has been crucial
to the development of mankind, providing both a regular
staple and a source of compounds essential for life carbohy-
drates, proteins, fat, and secondary metabolites. Its history as a
crop begins in the Fertile Crescent, a cradle area that today
includes part of Iran, Turkey, Syria, and Jordan. The domesti-
cation of its wild ancestor, with a two-row brittle rachis allow-
ing shattering of the seeds, Hordeum spontaneum (Figure 1),
which is still to be found in the Middle East, dates to the
Neolithic (10 000 years BC).
It is from here that the species set out first to colonize the
Mediterranean Basin and then, following the great river and sea
migration routes of the time, fanned out to Central and Northern
Europe, the Himalayas, and the Far East. Human intervention
has been a decisive factor in the breeding of this species down
through the millennia, the cultivars currently available manifest-
ing completely different traits from those found in either its
original parent or those populations that have evolved in the
wild. During the process of domestication, barley has gradually
accumulated traits that facilitated agricultural production. The
perfecting of crop management techniques during the period of
Roman dominion contributed to the spread of barley in the
fertile as well as in the poorer soils of the empire.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00021-8
The Mediterranean lands featured two barley types with
very precise traits. The first was the six-row type because it
grew rapidly in the mild Mediterranean climate, characterized
by low and erratic rainfall. The second, called two-row barley,
features high-quality grain and was sown in spring for direct
human consumption. Gladiators, whose diet consisted almost
entirely of food made from barley flour, were even called
hordearii, meaning barley men. Genotypes that had a naked
kernel were often resorted to as foodstuffs because of their
superior nutritional value. It was during the Roman era too
(Figure 2) that all the criteria for a rational use of grain and
straw were defined and refined to perfection, Columella pro-
viding a detailed description in his Res rustica. The yield per
hectare in the fertile lands was about tenfold the number of
seeds employed. Barley further expanded its horizons in con-
junction with Columbuss second voyage, when it was intro-
duced to the New World. Later, Dutch, English, French, and
Danish colonists promoted its spread throughout North Amer-
ica, as did their Spanish and Portuguese counterparts in Cen-
tral and South America. On the Altiplano of the Andean
nations of Peru and Bolivia, barley is grown for grain at higher
elevations over 4550 m than oat, wheat, and maize.
Morphology and Botany
Since the turn of the twentieth century, our insights into plant
growth, reproduction, and trait inheritance have been of indis-
pensable support to barley field management and the improve-
ment of cultivars. Several morphological forms have evolved,
including winter, spring, two-row, six-row, awned, awnless,
hooded, naked and covered, malting, feed (grain and forage),
and food type. The high diversity of barley phenotypes, true
diploid nature (2n 2x 14), and ease of hybridization and
cultivation have made barley a model genetic organism since
the rediscovery of Mendels laws. Barley belongs to Triticeae
and has seven pairs of distinct chromosomes, five of them
being without satellites (1H, 2H, 3H, 4H, and 7H) and two
with satellites (5H and 7H). Cultivated barley is an inbreeder
and it has more or less cleistogamous flowers, but under stress
conditions, it can be more open flowering. The normal out-
crossing rate has been determined to a maximum of 1.71.8%.
Despite a very high inbreeding rate, a high degree of diversity is
maintained, which is a sufficient basis for rapid adaptation to
varying environmental conditions.
Barley as Model for Triticeae Genetics and Genomics
Barley is a model organism for genetic and genomic studies in small
grain cereals because of several reasons, ranging from its high
degree of natural variation and wide adaptability to the diploid
147
148 THE CEREAL GRAINS | Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses

Figure 1 Fragile spike of Hordeum spontaneum.

Figure 2 Corona obsidionalis made with six-row barley spike as virtus

exercitus Romanorum.

genome structure and the strongly inbreeding-based mating

system, as well as the wide array of genomic resources. Its
seven chromosomes represent the basic genome of all Triticeae, Figure 3 Developmental mutants of barley spike.
and the first complete release of genome sequence has been
publicly available from October 2012, overcoming the difficul-
ties linked to the size (about 5500 Mb) and the presence of
repetitive DNA regions. The springboard for the sequencing of Neolithic*
Roman Renaissance
the barley genome has been offered by key molecular tools 500
period Pre-Mendel
developed in the near past, like barley ESTs (expressed seeds/m2
2000 seeds/m2 3000 seeds/m2
0.10.15
sequence tags), BACs (bacterial artificial chromosomes), and 0.40.5 t/ha 0.60.8 t/ha
t/ha
the physical maps.
Historically, barley genetic studies have their foundations
in Mendelian mutants, characterized by altered physiology Post- Today
and/or morphology and still preserved as Barley Genetic Stocks 1970
Mendel 20 000
14 000 seeds/m2
(Figure 3). In this regard, there are examples of morphological 8000 seeds/m2 seeds/m2
5 t/ha
mutations described in the past for which the gene/genes 2.5 t/ha 910 t/ha
responsible have been recently cloned and characterized. An
example is the Nud gene that controls lipid biosynthesis in the
pericarp, which glues the husks to the kernels and results in
either covered or naked barley seeds. Figure 4 Estimated yield potential of barley from domestication
Genetics knowledge has been applied to barley breeding, a (10 000 BC) to the beginning of twenty-first century. *Hordeum
dynamic process that has its pillars, on one hand, from short-, spontaneum in situ conservation.
medium- and long-term objectives and, on the other hand, on
perfecting selection criteria. Figure 4 traces the progress of genotyping platforms based on different types of molecular
barley production obtained in the course of history thanks to markers. The new genomic platforms based on NGS technolo-
successful breeding activities. gies are providing opportunities to advance radically in the
Structural and functional genomics, which merged in the inte- discovery of beneficial genes and alleles for barley breeding.
grative genomics, is supporting a knowledge-oriented breeding Functional genomics use transcriptome analysis that is, com-
to design the barley for the future in which good alleles will be prehensive and high-throughput analysis of gene expression for
operating all together in a superior genotype. Barley structural screening candidate genes, predicting gene function, discover-
genomics has developed EST sequences databases and ing cis-regulatory motifs, and characterizing transcriptional
 THE CEREAL GRAINS | Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses
regulatory networks models and reverse genetics. Genome-wide
reverse genetics have been developed in barley to understand the
genefunction relationships, including both transgenic strategies,
like insertional mutagenesis or activation tagging, and non-
transgenic ones, like TILLING (targeting induced local lesions in
genomes). Even metabolomics that is, the characterization of
metabolites present in a tissue in a specific environment
condition can be very informative. Integrative genomics deals
with combining the different data from these multiple-omics
technologies to study biological systems multilaterally. In partic-
ular, the combined analysis of a relevant amount of genotyping
and phenotyping data can give information useful for genetic
mapping and advanced breeding programs of barley. Barley
genetic resources remain the basis for this flowchart. For
this reason, barley collections representative of existing natural
genetic diversity are maintained worldwide in situ and ex situ,
in the gene banks. New barley genetic resources are now devel-
oped for more efficient phenotypinggenotyping, like nested
association mapping (NAM) populations obtained by crossing
diverse genotypes to a reference parent or multiparent advanced
generation inter-cross (MAGIC) based on the cross of several
parentals.
Talking about genetics, it is required to talk even of epige-
netics: epigenetic effects have been shown to be strongly
involved in developmental and physiological processes in
model plants. Although barley has been poorly investigated
from this point of view, some evidence exists on the impact of
epigenetic factors on seed development and response to
hormones.
Finally, from a practical point of view, what we need is not
new: efficient and inexpensive tools to select for agronomic
and quality traits. Molecular markers developed in the last
decades have been demonstrated to be of great help, in partic-
ular for pyramiding resistance genes in a marker-assisted selec-
tion (MAS) scheme of breeding. What is around the corner is
genomic assisted selection (GAS) that means to build breeding
schemes based on the selection of large portions of the
genome. This strategy has been applied successfully in barley
prebreeding/breeding for selecting barley genotypes improved
for disease resistance and other agronomic traits.
Barley Use
Today, 8090% of barley grain yield throughout the world is
earmarked for livestock feed, about 10% is slated as malt
production for beer, whisky, and other alcoholic beverages,
and only a very small fraction is still directly consumed by
man. Barley is still used today, albeit on a much reduced
scale, as a surrogate for coffee, under the guise of peeled barley
in the preparation of soups, and as an important staple in the
diet of populations in North Africa and Asia. Barley grain is
utilized in the diets of ruminants and nonruminant livestock,
poultry, and fish. Attempts to improve the kernel for livestock
feed have been focused on upgrading the nutritional value of
proteins, the digestible energy of meals, and gum content
(beta-glucan). A great deal of work has been done on proteins,
in that barley prolamins (hordein) are lacking in essential
amino acids, particularly lysine. However, five paths have
been identified to improve barley grain quality for animal
149
feeding: improving grain protein quality, reducing grain phytic
acid content, reducing beta-glucan content in feed for mono-
gastric animals, reducing rumen grain fermentation rate, and
increasing grain starch content. Barleys economic importance
also extends to the whole plant, which can be used for both
livestock feed and industrial purposes. The use of the whole
plant harvested at the end of the milk-ripe stage for silage is a
practice that is becoming increasingly common in a number of
barley-producing areas. This approach makes it possible to
develop the double-cropping system in irrigated fields by grow-
ing in the same soil and same year barley silage followed by a
crop of summer maize, sorghum, or soybean. Silage quality
can be improved only when varieties exhibit an overall grain/
biomass ratio (harvest index) greater than 50%.
As food, the human health benefit of consuming barley
grain in food formulations is well known. The kernel of barley
has a level of beta-glucans that is much higher than that found
in other grains. Beta-glucans are a kind of soluble dietary fiber
having hypoglycemic and hypocholesterolemic effects, as it
swells in the presence of water to form highly viscous solutions
that can trap glucose and cholesterol and slow down their
absorption. The reduction of the rate at which glucose is
released to the intestine and to the blood is particularly inter-
esting when barley flour is included in the preparation of
bread, a high glycemic food. Thus, incorporation of beta-
glucans derived from barley into this widely consumed food
can reduce the glycemic response to its consumption. The use
of beta-glucan-enriched barley flour is recommended to pro-
duce bread dough made of wheat/barley flour blends with a
not-too-high proportion of barley flour. The majority of the
Waxy barley genotypes (which have a very low to null amylose
content) also have high beta-glucan content and can therefore
be used to improve the hypocholesterolemic effects of foods.
Barley flour can be easily incorporated into many wheat-
based baked or not baked products such as cookies, bread,
flatbread, tortillas, puffed sacks, and pasta for producing the
so-called functional foods (Figure 5). In addition, tocols and
phenolic compounds of barley are also considered bioactive
components.
Figure 5 Functional pasta type made with durum wheat semolina
(75%) and barley flour (25%).
150 THE CEREAL GRAINS | Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses

Malting Quality during mashing. During the malting process, high production
and/or activation of hydrolytic enzymes is required for cell-
Barley is, and has been since the dawn of civilization, the cereal
wall degradation and protein solubilization, but starch break-
par excellence in the production of malt for beer, whisky, and
down should be minimal, to avoid unwanted consumption of
other alcoholic beverages. Indeed, ever since the development
utilizable dry matter by the embryo. To this aim, malting
of the most essential techniques by the ancient Egyptians and
should both accelerate modification and retard embryo
Babylonians, the art of brewing has continued to be perfected
growth, which are partially conflicting purposes.
down through the ages and its secrets jealously guarded by the
It is well known that the higher the level of protein in the
master brewers, despite the fact that fermentation technology
barley grain, the lower the percentage of carbohydrate and,
had remained entirely shrouded in the dark until the nine-
then, the extract yield of the malt. In addition to this negative
teenth century.
relationship with starch level, storage proteins have been sug-
The transformation of barley into malt and of the latter into
gested to play a significant part in determining the rate at
beer is a biotechnological process that is markedly dependent
which the whole endosperm is broken down during malting.
on the initial raw material, that is, on the barley grain that gives
More specifically, malting barley must have soft and mealy
rise to the malt. To this aim, barley has first to be converted
rather than hard and flinty endosperm and include such other
into malt, to increase its content of enzymes. Among them,
characteristics as high germinative energy, a certain degree of
amylases are needed to convert polymeric starch into soluble
dormancy to prevent pregermination in case of rain at harvest,
sugars, which can be used by yeasts for alcoholic fermentation.
adequate enzyme activity (amylase, beta-glucanase, and prote-
Malting is a controlled process of germination. As the first
ase), low beta-glucan content, low protein content, high starch
step of malting, barley is steeped to imbibe the grains and start
content, reduced glumeal hulls, and high extract percentage.
germination. After 13 days of germination, depending on the
The percentage of extract is directly linked to starch content
malting conditions, the aleurone layer responds to gibberellin
and seed size and inversely correlated to protein content. A
hormone from the embryo and releases hydrolytic enzymes
high nitrogen content adversely affects malt production by
that disaggregate and solubilize the endosperm. They can break
reducing the amount of extract and lowering quality. This is
down starch and storage proteins into simple molecules that
because a longer period is required for the modification pro-
nourish the growing embryo. Since starch and storage proteins
cess, which results in a greater development of rootlets and
are enclosed in the walls of the dead endosperm cells, these cell
coleoptiles and consequent respiratory losses, and because of
walls have to be dismantled to allow access of the hydrolytic
an abundance of soluble proteins that enter the extract and
enzymes to the internal reserves. Breakdown of endosperm cell
form suspensions in the beer. It is, however, important that the
walls mainly involves degradation of beta-glucans, which are
extract contain a permanent fraction of soluble nitrogen
the major constituent of the walls (>70%). This initial step is
(0.5%) to feed the yeasts during fermentation.
performed by endo-(1!3),(1!4)-beta-D-glucanase (shortly,
Malting begins with the soaking of the kernel in water to
beta-glucanase), which is the chief enzyme responsible for
raise its moisture content to 4245%, followed by the period
beta-glucan degradation and is therefore the enzyme leading
(4 days) of modification. The changeover from barley to malt
endosperm modification. Modification is a term used to
commences when the embryo secretes gibberellic acid, which
mean all the chemical and physical changes that occur when
immediately stimulates the production of enzymes (glucanase
barley is converted into malt, particularly the weakening in the
and pentosanase) that alter the endosperms structure by dis-
consistence of the endosperm consequent to the degradation
solving the walls of its cells and freeing starch grains. Other
of the cell walls. Prior to milling, malt appearance resembles
enzymes as phosphatase, phytase, glucanases, pentosanases,
the original barley grain but is friable when crushed, revealing
and protease also become active in the early stages of modifi-
the physicalchemical changes that have occurred during malt-
cation, while the amylases are triggered later. All of this activity,
ing. Secretion of beta-glucanase and other hydrolytic enzymes
which serves as a vehicle to move substances from the endo-
starts from the embryos scutellum and proceeds in the aleu-
sperm to the embryo for the formation of new tissues and
rone layer from near the embryo to the distal end of the grain.
organs (rootlet and coleoptile), is accompanied by a marked
Much research has focused on the role of endosperm compo-
increase in respiration: the resulting losses can range from 5%
nents in determining these aspects, and beta-glucans are con-
to 9% unless restricted by a rapid and uniform germination.
sidered to play a negative role. On the other hand, beta-glucans
From what we have mentioned thus far, it is not difficult to
have also received much interest recently with regard to poten-
see that malting quality depends on a host of organic factors.
tial health benefits. Thus, whereas high beta-glucans are
The analytic data of a single character are insufficient to judge
required for food functionality, low beta-glucans are usually
either the grain quality initially or the malt quality thereafter.
desirable for the best malting quality.
Genetically speaking, breeding for malt varieties is a com-
The endosperm of unmalted grains is compact and inacces-
plex issue because many traits at grain, malt, and wort level
sible to hydrolytic enzymes because large proteins cannot per-
have to be taken into account.
meate cell walls. Thus, in the germinating barley grain,
thorough breakdown of endosperm cell walls and partial
digestion of the protein matrix inside the endosperm cells are Biotic and Abiotic Stress Resistance
necessary steps to render starch granules accessible to amylo-
lytic enzymes. These two processes are the chief causes of the Barley, despite its proven adaptability to a range of ecological
physicalchemical modification of the endosperm that occurs situations throughout its evolution, is nonetheless susceptible
during malting and make the starch susceptible to degradation to extreme environmental situations and a number of
 THE CEREAL GRAINS | Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses
Diseases
Heat shock
Lodging
Cold
Soil of low fertility
Salt Anoxia and drought
Heavy metals Ions - acid and alkaline
Figure 6 Traits to cope with biotic and abiotic stresses in barley.
pathogens (Figure 6). These adversities can determine crop
stress, ensuing yield losses of varying magnitude. Such field
management practices as irrigation, fertilization, and pest con-
trol provide a certain measure of crop protection, although
they increase the energy input required for successful harvests.
Studies of genetic resistance have focused intensely on the
selection of those plant genotypes less subject than others to
the insults of a biotic and abiotic nature so as to increase yields
and yield stability.
Abiotic Stress Response
The optimum development of barley over its life cycle depends
on a number of environmental stress factors that can prevent
the plants expressing its maximum genetic potential. Severe
grain losses are often caused by high or low temperatures,
drought, anaerobiosis, and such soil anomalies as excess salt
or heavy metals. The responses elicited from the plant by these
stresses, when not lethal, include alterations in its processes of
photosynthesis, respiration, and hormonal regulation through
the development of specific, adaptive defense systems and
mechanisms that are molecularly controlled. The duration of
the stress and the plants growth stage at the formers onset in
turn affect yield. One can also find differing reactions as to
plant susceptibility to adverse conditions. Thus, genetic vari-
ability plays a primary role in determining positive adaptation
to environmental stresses and, hence, in supporting the spread
of various barley genotypes to extreme climatic conditions. The
integration of molecular genetics with physiology is leading to
the identification of the most relevant genes controlling abiotic
stress tolerance with the final aim to develop high yielding
cultivars able to improve performance in both high- and low-
yielding environments.
Biotic Stresses
The Compendium of Barley Diseases lists 37 viral, 36 fungal, and 4
bacterial diseases, including 6 nematodes as well as a
151
substantial number of insect species, which regularly or casu-
ally cause damage during the entire life cycle. The level of crop
losses in barley at global scale, due to diseases and pests, has
been estimated to be 10.5% and 8.5%, respectively. The fight
between barley and its parasites is a dynamic process in which
two different strategies of resistance are activated. The first type
is resistance to infection synonymous of hypersensitivity, race-
specific, vertical, or major gene resistance. This type works as
gene-to-gene interaction between barley genes for resistance
and pathogen genes for avirulence. The second type is defined
as quantitative or partial resistance, field resistance, durable
resistance, non-race-specific resistance, horizontal resistance,
or minor gene resistance. The diseases and insects inflicting
severe crop losses on a worldwide scale are described here
(Table 1).
With the development of molecular marker technology,
marker-assisted selection (MAS) has been routinely used as
tool for detecting genetic loci underlying the traits for resis-
tance and for incorporating them in elite genotypes. Although
chemical treatments and agronomic techniques are effective in
controlling pests and diseases of barley, the availability of
resistant cultivars is the most economic and environmentally
sound strategy for reducing losses.
Conclusions and Future Prospects
Barley today ranks fourth behind wheat, rice, and maize
among the worlds cereals for the importance of its contribu-
tion, whether direct or indirect, to the production of food. The
global production is estimated in 2013 as 135 millions of
metric tons (Mt) in a harvested area of 50 million hectares
(Mha) with an average of 2.7 metric tons (t ha1) of grain per
hectare. Grown in almost every country throughout the globe,
it is generally found as a crop in areas that are less conducive to
the survival of other cereals. It is very widespread in the
drought-prone lands of the Near East, Northern Africa, and
the highlands of India; in the saline soils of Australia; in Nordic
countries; and in areas that, while fertile, are marked by a very
short growing season (<100 days).
This flexibility is doubtlessly due to barleys range of genetic
variants. The plant appears capable of producing yields that are
competitive with yet require less energy input than other
cereals. Barley as a winter crop can register yields as high as
10 t ha1 of grain at lower costs than the other cereals due
mainly to high tillering capacity and its high nitrogen use
efficiency (NUE) and water use efficiency (WUE). Barleys
capacity to produce biomass even under restrictive conditions
implies that in all likelihood, it will continue to play a strategic
role in food production for the foreseeable future in many areas
of the world where drought and other adverse climatic phe-
nomena are inimical to the plant growth and development.
Due to phenotypic plasticity, further improvements in yield
potential and stability can be expected in the near future in the
modern barley farming system.
Exercises for Revision
Are specific barley varieties used for malting?
Explain the difference between naked and hulled grains.
152 THE CEREAL GRAINS | Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses

Table 1 Major diseases and pests of barley Are the barley mutants useful to increase secondary metab-
olites for food?
Disease common name Causal agent
Which strategy is still active for disease resistance in barley:
Fungal disease monogenic or quantitative resistance?
Barley stripe Pyrenophora graminea
Powdery mildew Blumeria graminis
Net blotch Pyrenophora teres See also: Agronomy of Grain Growing: Barley: Agronomy;
Fusarium Head Blight Fusaria
Barley, Rice and Maize Processing: Barley: Malting; Barley:
Snow rot Typhula incarnata
Spot blotch Cochliobolus sativum
Milling and Processing; Beverages from Grains: Beverages:
Leaf blotch or scald Rynchosporium secalis Distilled; Fermentation: Foods and Nonalcoholic Beverages;
Loose smut Ustilago nuda Fermentation: Origins and Applications; Bioactives and Toxins:
Ramularia leaf spot Ramularia collo-cygni Bioactives: Antioxidants; Mycotoxins; Breeding of Grains: Barley:
Yellow rust Puccinia striiformis Genetics and Breeding; Carbohydrates: Health Effects of b-Glucans
Brown rust Puccinia hordei Found in Cereals; Cereals: Chemistry and Physicochemistry of Non-
Black stem rust (including Puccinia graminis starchy Polysaccharides; Glycemic Index, Glycemic Load, and Their
Ug99) Health Benefits; Resistant Starch and Health; Food Grains and Well-
Septoria leaf blotch Septoria passerini being: Functional Foods: Overview; Genetics of Grains: Genomics
Rhizoctonia root rot Pythium
of Food Grains; Grain Harvest, Storage and Transport: Barley:
Root rot Rhizoctonia solani
Ergot Claviceps purpurea
Harvesting, Storage, and Transport; Grain Marketing and Grading:
Covered smut Ustilago hordei Barley: Grading and Marketing; Non-food Products from Grains:
Loose smut Ustilago nuda Cereal Grains as Animal Feed; Processing of Grains: b-Glucans:
Take-all Gaeumannomyces graminis Measurement and Processing; The Basics: The Grain Crops: An
Eye-spot Pseudocercosporella herpotrichoides Overview; The Cereal Grains: An Overview of the Family of Cereal
Leaf non parasitic spots Unknown Grains Prominent in World Agriculture; Cereals: Domestication of the
Bacterial disease Cereal Grains.
Black chaff Xanthomonas translucens
Bacterial blight Xanthomonas campestris
Viral disease
Barley yellow mosaic virus- Potyviridae (soil borne disease
BAYMV, BAMMV transmitted by Polymixa graminis) Further Reading
Barley yellow dwarf virus Luteoviridae (transmitted by aphids)
BYDV Bulgarelli D, Biselli C, Collins NC, et al. (2010) The CC-NB-LRR-type Rdg2a resistance
Pests and nematodes gene confers immunity to the seed-borne barley leaf stripe pathogen in the absence
Cereal cyst nematode Heterodera avenae of hypersensitive cell death. PLoS One 5(9): e12599.
Demetriou K, Kapazoglou A, Tondelli A, et al. (2009) Epigenetic chromatin modifiers in
Root lesion nematode Pratylenchus spp.
barley: I. Cloning, mapping and expression analysis of the plant specific HD2 family
Green bug Elymus canadensis of histone deacetylases from barley, during seed development and after hormonal
Aphids treatment. Physiologia Plantarum 136: 358368.
Hessian fly Mayetiola destructor Fincher GB (2011) Biochemistry, physiology, and genetics of endosperm mobilization
Cereal leaf beetle Oulema melanopus in germinated barley. In: Ullrich S (ed.) Barley: Production, Improvement and Uses,
Cereal ground beetle Zabrus tenebroides pp. 449477. Chichester, UK: Wiley-Blackwell.
Frit fly Oscinella frit Finocchiaro F, Ferrari B, Gianinetti A, et al. (2012) Effects of barley beta-glucan-
Cutworms Agrotis enriched flour fractions on the glycaemic index of bread. International Journal of
Food Sciences and Nutrition 63(1): 2329.
Francia E, Tondelli A, Rizza F, et al. (2011) Determinants of barley grain yield in a wide
range of Mediterranean environments. Field Crops Research 120: 169178.
Is the whole barley plant used for forage? Graner A, Kilian A, and Kleinhofs A (2011) Barley genome organization, mapping and
Explain the difference between winter, alternative, and synteny. In: Ullrich S (ed.) Barley: Production, Improvement and Uses, pp. 6384.
Chichester, UK: Wiley-Blackwell.
spring barley varieties. Mathre DE (1997) Compendium of Barley Diseases. St. Paul, MN: American
Are there in barley grain secondary metabolites for human Phytopathological Society.
health other than beta-glucans? Paulitz TC and Steffenson BJ (2011) Biotic stress in barley: Disease problems and
Why is the barley plant considered more economically and solutions. In: Ullrich S (ed.) Barley: Production, Improvement and Uses,
pp. 307354. Chichester, UK: Wiley-Blackwell.
environmentally suitable than other cereal crops? Rizza F, Pagani D, Gut M, et al. (2011) Diversity in the response to low temperature in
What is the barley genome size? representative barley genotypes cultivated in Europe. Crop Science 51: 27592779.
Stanca AM, Tumino G, Pagani D, et al. (2013) The Italian barley genetics mutant
collection: conservation, development of new mutants and use. Advance in Barley
Sciences 3035.
Exercises for Readers to Explore the Topic Further Thomas WTB, Hayes P, and Dahleen L (2011) Application of molecular genetics and
transformation to barley improvement. In: Ullrich S (ed.) Barley: Production,
WUE (water use efficiency) and NUE (nitrogen use Improvement and Uses, pp. 122143. Chichester, UK: Wiley-Blackwell.
efficiency) are considered two strategic traits to develop Ullrich S (2011) Significance, adaptation, production and trade of barley. In: Ullrich S
plants to cope with environments. Is barley a more efficient (ed.) Barley: Production, Improvement and Uses, pp. 313. Chichester,
UK: Wiley-Blackwell.
species within cereals? von Bothmer R and Komatsuda T (2011) Barley origin and related species. In: Ullrich S
How do the agronomic practices improve yield and quality (ed.) Barley: Production, Improvement and Uses, pp. 1462. Chichester,
of the barley grain? UK: Wiley-Blackwell.
 Sorghum Grain, Its Production and Uses: Overview
A Cruickshank, DAFF Queensland, Warwick, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The sorghum grain is a high-energy cereal grain that is a
staple to many poorer farmers in Africa and parts of Asia.
Sorghum is a genetically diverse and robust plant, posses-
sing resistances to many biotic and abiotic stresses.
Sorghum is a (sub)tropical grass (like sugarcane and maize)
with a C4 photosynthetic pathway.
Sorghum bicolor is classified into five races: bicolor, kafir,
caudatum, durra, and guinea.
Major sorghum producers are Nigeria, the United States,
India, Argentina, Ethiopia, and Australia.
Sorghum can grow in temperatures from 15 to over 40  C
Flowering and seed set require temperatures between 20
and 39  C.
Significant diseases are downy mildew, ergot, and grain
mold.
Significant insect pests include sorghum midge and
greenbugs (aphids).
Sorghum grain can be white, yellow, red, or purple.
Sorghum is consumed as a thin porridge or gruel, as a thick
porridge or dough, as various flatbreads, as popped or
puffed sorghum grain, or as alcoholic beverages.
Learning Objective
To achieve an overall understanding of sorghum as a grain
worldwide: its origins, breeding, agronomy, production,
processing, and utilization
Introduction
All grain sorghum is the product of cultivated types of Sorghum
bicolor (L.) Moench. Sorghum bicolor is an African species and
the crop was first domesticated in East Africa. In traditional
agriculture, tall varieties of sorghum are grown for multiple
purposes including various uses of grain, fodder, and extract of
sweet syrup (by crushing stems in the same manner as with
sugar cane) and use of the panicles for broom heads. Today,
sorghum is a staple for approximately half a billion people.
In developed agriculture, particularly in the Americas and
Australia, sorghum has been selected and bred for specific
end uses grain sorghums as short plants suited to mechani-
zed agriculture and taller types for forage, fodder, and silage
production.
Grain sorghum is a crop with great genetic diversity and
adaptation to drier and hotter environments. It has tradition-
ally been used in many different ways in Africa and Asia. It is
now an important world crop and the most important crop
locally in some hot and dry regions.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00023-1
The Origins of Domesticated Sorghum
Botanically sorghum belongs to the grass family and the tribe
Andropogoneae. This is a tropical and subtropical group of
grasses, which also includes sugarcane (Saccharum spp.) and
maize (Zea mays). Like all the grasses in this tribe, sorghum has
a C4 photosynthetic pathway. The genus Sorghum has at least
20 species extending from Africa through Asia and including
12 species endemic to Oceania. Sorghum bicolor, a native of
Africa that has spread throughout the world, has three subspe-
cies: S. bicolor bicolor, S. bicolor verticilliflorum, and S. bicolor
drummondii, with distinct morphology for the respective
heads (Figure 1).
S. bicolor subsp. verticilliflorum is the wild progenitor spe-
cies. These sorghums have small seed, < 1 mm thick. These
seeds are shed soon after they ripen. This process is called
shattering, and selection of nonshattering off-types is one
of the first steps in domesticating a wild plant to develop
a crop.
S. bicolor subsp. bicolor incorporates all the cultivated types.
The cultivated types have been further classified on the
basis of the shape of heads and grain into five races: bicolor,
kafir, caudatum, durra, and guinea. The range of head types
is illustrated in Figure 2.
S. bicolor nothosubsp. drummondii incorporates all material
derived from natural crossing between S. bicolor subsp.
bicolor and S. bicolor subsp. verticilliflorum types. In some
production systems, weedy S. bicolor nothosubsp. drum-
mondii types (known as shattercanes) are problem weeds.
Analysis of DNA sequence data has led scientists to believe
there have been at least two, possibly more, domestications
of sorghum. There is little doubt that the original domestica-
tion occurred in East Africa somewhere in Ethiopia or possi-
bly Sudan. Evidence from DNA sequence analysis suggests that
there has been at least one other more recent domestication in
West Africa, giving rise to the group that has been distinguished
by the name margaritiferum. These plants have a lot of wild
plant features and are grown separately from other sorghums,
named as a different crop and prepared in different foods.
Sorghum Production
Sorghum is produced in the tropics and in subtropical or
temperate regions throughout Africa, Asia, and the Americas
(only a small amount of grain sorghum is produced in
Europe). Prior to the colonial era, maize was the dominant
grain crop in the Americas and sorghum the most important
summer cereal in Africa. Since both crops have been available
worldwide, there has been a general trend for maize to be more
popular in higher rainfall areas and sorghum to be cultivated
153
154 THE CEREAL GRAINS | Sorghum Grain, Its Production and Uses: Overview
Figure 1 Heads of the three subspecies of Sorghum bicolor: left S.
bicolor subsp. verticilliflorum, middle S. bicolor nothosubsp.
drummondii, and right S. bicolor subsp. bicolor.
Figure 2 Variation in head type for Sorghum bicolor.
in more marginal environments. Some exceptions to this occur
where either sorghum or maize is more suited to preparation
of an important food. Africa produced 40% of the worlds
sorghum in the 10 years from 2004 to 2013, and West Africa
produced just over half of that. Over the same period, 37% of
world sorghum production was in the Americas and 20% in
Asia and Oceania (Figure 3).
Sorghum is produced in over 100 countries, but of these,
only 14 produce about a million tonnes or more annually (mt
pa) (Table 1). During the decade 200413, the United States
produced the most sorghum (averaging just under 10 mpa) at
an average yield of 4 t ha 1. However, there has been a down-
ward production trend in the United States over the past 50
years. Other major sorghum producers are Nigeria, India, and
Mexico, each producing 68 mt pa in the decade 200413.
Nigeria experienced dramatic growth in sorghum produc-
tion from 1964 to 2006 with a slight decline since then.
Ethiopia is also a major producer of sorghum in Africa with
a marked increase in sorghum production since 2000. The
Ethiopian increase has been characterized by average yields
from 2004 to 2013 of 1.8 t ha 1 compared to an Africa-wide
average of <1 t ha 1. Sudan, Burkina Faso, Niger, Mali, and
Cameroon have all steadily increased their sorghum produc-
tion and currently average 11.5 mt pa. Sudan has the highest
ratio of sorghum to maize in the world, this being an indicator
of the importance of sorghum to a particular country.
India is still a major sorghum producer, but there has been
a decline of production since 1995, due to a number of chang-
ing factors including the increased capacity to irrigate higher
value crops such as rice, and the cost and availability of labor
for sorghum production. In the 1960s, China produced over
8 mt of sorghum pa, but this has declined to around 2 mt pa.
Over this time, sorghum yields in China have improved to an
average of just under 4 t ha 1. Total production appears to
have declined in China, apparently for similar reasons to the
changes in India, particularly involving the greater availability
of rice. Consumption of sorghum as food in China has
declined but the use in distilled spirits has increased.
Mexican sorghum production has increased over the past 50
years but appears to have stabilized at just over 6 mt from 2007
to 2013, with impressive average yields of 3.7 t ha 1. Mexico
consumes all the sorghum it can produce plus significant
imports from the United States. In the early 1980s, Argentina
produced approximately 8 mt pa but this has since fallen dra-
matically. Despite sorghum being displaced by herbicide-
resistant maize and soybean, the average yield of sorghum in
Argentina is over 4.5 t ha 1. Brazil produced 1.8 mt pa at a yield
of 2.4 t ha 1 over 200413, the result of steady increase over the
past 50 years.
Australia is the only non-African country where sorghum
production significantly exceeds that of maize. A study using
simulation to compare climate-adjusted yield improvement
found sorghum yield improvement was substantially better
than that of wheat in subtropical Australia. This has contrib-
uted to sorghum production rising to average 2.1 mt pa and
yields of 3.1 t ha 1 over 200413.
Adaptation of the Sorghum Crop
Although it is essentially a tropical and subtropical species, the
cultivated form of sorghum is also grown to 40 latitude as a
warm season crop. Generally, sorghum needs soil temperatures
over 15  C for crop establishment. Crop growth can occur from
these temperatures to just over 40  C. Flowering and seed set
require temperatures between 20 and 39  C, the most sensitive
process being pollen production with higher and lower tem-
peratures leading to sterility. Recent research has found genetic
variation in the extreme temperature tolerances of both germi-
nation and flowering. This means there is the capacity to
improve grain sorghums tolerance of extreme weather events
that are projected to increase in the future. Sorghum as a species
also possesses many characteristics that enable growth and
development in water-limited environments including stay-
green drought resistance during grain-filling and transpiration
efficiency. Many traditional landrace varieties are sensitive to
photoperiod, requiring days shorter than a certain maximum
before they will flower and produce grain. Such varieties may
fail to flower and produce grain when grown at different
 THE CEREAL GRAINS | Sorghum Grain, Its Production and Uses: Overview 155

Western Central
South-Eastern Asia Oceania
Asia Asia
Middle Africa Eastern Africa

Eastern Africa Middle Africa

Northern Africa

Southern Asia Western Africa

Eastern Asia Southern Africa

Northern Africa
Europe
Northern America
South America
Central America Caribbean
Western Africa Central America
Eastern Asia
Southern Asia
Caribbean South America
South-Eastern Asia
Western Asia
Central Asia
Northern America Southern Africa
Oceania
Europe
Figure 3 Sorghum production worldwide by region, averaged for the decade 200413. Source: FAOSTAT.

Table 1 Production of sorghum by major countries, averaged for the the soil, but fertilization with phosphorus, potassium, and
decade 200413 other minor nutrients will often still be necessary to remove
limitations to yield.
Country Tonnes % The agronomy of sorghum production in mechanized agri-
culture is quite different to that in traditional production sys-
World 59 850 648 100
The United States 9 327 805 15.6 tems. In developed agriculture, sorghum is grown as a row crop
Nigeria 7 891 520 13.2 at plant densities suited to the available moisture. Nitrogen
India 6 883 880 11.5 fertilizer may be applied entirely preplanting or split between a
Mexico 6 361 639 10.6 starter application and a later targeted top-up when the likely
Sudan (former) 3 873 500 6.5 yield is more predictable. Herbicide application may be used
Argentina 3 056 573 5.1 to desiccate the sorghum plant and grain in preparation for
Ethiopia 2 943 216 4.9 harvest.
Australia 2 171 743 3.6
China, mainland 2 141 650 3.6
Brazil 1 813 631 3.0
Burkina Faso 1 673 123 2.8
Niger 1 018 760 1.7
Sorghum Breeding
Mali 993 038 1.7
Cameroon 961 818 1.6 Traditional farmers have been responsible for the selection and
maintenance of the great diversity that exists in the landraces of
Source: FAOSTAT. sorghum. When some of these landraces were introduced to
the Americas, any off-types that had dwarfing genes or different
maturities were selected and (just as important) were studied.
latitudes or even similar latitudes with different temperature An understanding of the differences in height and maturity
regimes. developed that allowed selection of types that were more suited
Sorghum is adapted to a wide range of soil conditions. to mechanized agriculture. Subsequently in the United States,
Some of the genetic characteristics that confer this adaptation there was a great plant breeding initiative called the sorghum
are being discovered: a general tolerance of salinity, greater conversion program in which landraces were converted by
tolerance to acidic aluminum ions, more efficient uptake of crossing them to short, photoperiod-insensitive types and the
essential phosphorus, and different angles of nodal root progeny back-crossed to the landrace while selecting for short
growth. As with other cereal grains, in most circumstances, photoperiod-insensitive plant type. This provided a resource to
nitrogen will be the limiting nutrient in grain production. sorghum breeders from which many improved traits could be
Sorghum will form mycorrhizal associations with fungi that bred into other genetic backgrounds including high yield,
improve the nutrient efficiencies of elements that are present in drought resistance traits, and midge resistance.
156 THE CEREAL GRAINS | Sorghum Grain, Its Production and Uses: Overview
Sorghum now has a published genome sequence that
allows breeders to develop understanding of the genetics of
traits of which their predecessors could only dream. Given the
great diversity available within the species, this understanding
should allow development of grain sorghum with robust adap-
tation to extreme environments and grain qualities that pro-
vide a range of products fit for many purposes.
Sorghum Diseases and Pests
Like all crops, sorghum productivity is affected by a range of
pests and diseases. Some pests or diseases have a specific inter-
action with sorghum; others are more widely adapted and are a
threat to other crops also. Some directly affect yield of usable
grain through killing plants or reducing the amount of grain
produced per plant. Diseases also indirectly influence produc-
tivity through damaging and devaluing grain or by causing
farmers to sacrifice potential yield with management decisions
that are part of a strategy to avoid disease.
To show the diversity of pest and disease biology, three
diseases, downy mildew, ergot, and grain mold, and two impor-
tant insect pests, sorghum midge and greenbugs (aphids), will
be considered.
Downy mildew is caused by the pseudofungus Peronoscler-
ospora sorghi. This pathogen can infect other species including
maize. It can survive for years in the soil as a resistant oospore
and can do most damage when oospores infect a young crop in
moist soils. This is called systemic infection and the pathogen
can then grow throughout the plant. Symptoms include ini-
tially leaf yellowing but eventually leaves are completely dead
and visibly shredded. When warm humid conditions favor
growth of the pathogen, it can produce short-lived spores
called conidia that can infect the aerial plant parts. Conidial
infection may cause minimal damage in mature plant or lead
to systemic infection in a very young plant. Control is achieved
by host plant resistance, seed treatment with fungicides, and
cultural measures such as crop rotation (to reduce oospore
infection) and early sowing (to reduce conidial infection
from plants infected earlier).
Sorghum ergot is caused by the fungus Claviceps africana (in
the Indian subcontinent by C. sorghi). It is host-specific to
plants of the genus Sorghum and has one very specialized
mode of infection: conidia of sorghum ergot can only infect
nonfertilized stigmas of flowering sorghum heads. Genetic and
environmental factors can contribute to nonfertilized stigmas
being exposed to infection. Male-sterile lines used in the pro-
duction of hybrid seed are particularly vulnerable; and seed
production is done in lower risk environments and protected
with fungicides if needed. The environmental factors that lead
to ergot disease are factors that prevent successful pollination:
rainfall and high humidity at flowering and cool temperatures
prior to and at flowering. The sensitivity of the flowering
period influences farmer decisions on when to sow sorghum,
often choosing not to sow sorghum after a certain date to avoid
a high risk of ergot infection.
Sorghum grains are exposed to the weather, and moisture
on the grain can lead to infection by a number of species of
fungi particularly Fusarium and Curvularia. Collectively, these
infections are referred to as grain mold. Some cause nothing
more than discoloration of the grain, but other species in some
circumstances produce toxins that make the grain unsuitable
for consumption. Many traditional varieties, particularly in
West Africa, flower in response to changes in day length so
that most if not all of their grain-filling period occurs after the
end of the rainy season.
The sorghum midge (Stenodiplosis sorghicola) is an insect
that is totally dependent on sorghum plants for its survival
and reproduction. The female midge seeks out flowering sor-
ghum to lay its eggs at the base of florets. The larvae of the
midge then feed on the ovary and prevent the development of
grain. In susceptible sorghum types, if the environment is
warm and humid, losses can be substantial. The level of dam-
age is also dependent on the number of flying female midge
present when a crop is flowering. Most commonly, these
females come from wild or volunteer sorghum (particularly
Sorghum halepense) or an earlier crop of cultivated sorghum.
Pupae can overwinter in plant trash in a diapause state. There
are sorghum lines that resist the oviposition or the larval
feeding. With resistant cultivars and cultural practices, such as
early sowing, good control of midge damage can be achieved
without the use of insecticide. In Australia, the widespread use
of midge-resistant sorghum has allowed the survival of many
more beneficial arthropods (predatory insects, spiders, etc.),
and this has aided integrated pest management in other sum-
mer crops also.
A range of aphid species will feed on sorghum but the most
common pest is the greenbug (Schizaphis spp.). They are grass
feeders and are also pests of other cereals in farming systems. In
sorghum, they cause damage by degrading tissue with their
saliva, by depletion of nutrients and energy, by contaminating
harvested grain with honeydew, and by being a vector for plant
pathogenic viruses notably maize dwarf mosaic.
Greenbugs are capable of both sexual reproduction and
parthenogenesis. They also have a short generation time and
a capacity for rapid population increase. This has led to rapid
emergence of new biotypes with different preferences in feed-
ing or resistances to pesticide, when control measures provided
selection pressure on the greenbug population. Resistant vari-
eties and natural enemies, together with careful use of
pesticide, can provide adequate control.
Sorghum Grain Composition and Utilization
The sorghum grain is a typical grass seed but it has been
selected over time for larger seed size and other particular
characteristics. Each grain is a caryopsis the botanical name
for a fruit containing a single seed with the fruit wall (pericarp)
and seed coat (testa) fused. The color of the grain is mostly
determined by the pericarp that can be white, yellow, red, or
purple (almost black), but the presence of condensed tannins
in the testa causes a strong brown subcoat. Tannin-rich sor-
ghums offer the advantage of being bird-proof to a degree, but
they also offer greater difficulty in most forms of processing.
Variations in head shape and grain color are illustrated in
Figure 4.
The embryo of a cultivated sorghum grain is larger than the
embryo of wild sorghum, but the increase is not proportional
to the increase in grain size with domestication. Most of the
 THE CEREAL GRAINS | Sorghum Grain, Its Production and Uses: Overview
Figure 4 A range of sorghum grain colors, including some white grains
stained by grain mold.
volume of the grain is made up of endosperm, but different
sorghum seed lines have very different endosperm structures
and textures, which determine their value for different uses of
the grain.
When the grain is mature, its endosperm is composed of
starch granules, spherical bodies composed of kafirin proteins,
and a protein matrix of glutelin. These three components can
be loosely packed in what is called floury endosperm or tightly
packed in what is called corneous endosperm. Kafirins can
form disulfide bonds, with one another and with glutelin,
which are very resistant to digestion. The resistance of these
bonds to hydrolysis is believed to reduce the digestibility of
both protein and starch, particularly in tightly packed corneous
endosperm.
There are other factors present in whole grain that can cause
resistance to digestion condensed tannins in the testa and
phytate in both the embryo and the pericarp. The tannins,
when present, can interfere with protein digestion. Phytate
can interfere with the digestibility of protein, phosphorus,
and iron.
The end uses of sorghum can be divided for convenience
into traditional uses associated with cultivation of landraces in
traditional African and Asian societies and uses in the devel-
oped economies of the Americas, Australia, and Europe.
Throughout Africa, sorghum grain is prepared as a wide
range of foods. Traditionally, the milling of sorghum grain
has been a laborious manual task with tools that are variations
on the theme of a mortar and pestle. The milled grain can then
be used in a variety of ways; broadly, most foods can be classed
as a thin porridge or gruel, as a thick porridge or dough, as
various flatbreads, as popped or puffed sorghum grain, or as
alcoholic beverages. In West Africa, the margaritifera types are
cultivated separately and often used in a manner similar to rice.
The variety in the foods produced is influenced by the choice of
sorghum, that is, use of the whole grain meal or some fraction
or use of a white-, red-, or brown-seeded sorghum containing
condensed tannins. Variation within these foods is also
157
Figure 5 Injera, a fermented flatbread popular in Ethiopia, made from
teff or sorghum. Source: Illustration from Wikipedia.
produced by the absence or the presence (and nature) of
fermentation. When sorghum is prepared as a heavy porridge,
it is usually eaten by hand with a savory sauce.
In Ethiopia, sorghum is made into injera (Figure 5), a
fermented flatbread that looks to western eyes like a pancake.
The preparation of injera includes a lactic acid fermentation
leading to a marked sour taste. The most valued grain for injera
is the Ethiopian grain teff, but sorghum is produced in much
greater quantities than teff and is preferred over wheat or barley
for making injera. Consequently, Ethiopia is one of the few
places in the world where sorghum attracts a higher price than
wheat. Injera is usually made with little or no bran. In Sudan, a
similar bread called kisra is made with whole grain meal.
In India, sorghum is used in roti-style flatbreads or as a
boiled grain substituting for rice. Rotis are often made with a
mixture with wheat flour. As in many other uses, the elastic
nature of the gluten in wheat flour helps maintain the structure
of a roti for longer than if made with sorghum flour only.
In East Asia, particularly China, sorghum grain is used
(either on its own or in combination with other grains) in
the production of highly alcoholic distilled spirits. These spirits
are produced for both high-value exclusive markets and more
traditional local consumption. Because the sorghum produc-
tion in China has declined and there is a demand for sorghum-
derived spirits in the growing middle class, China may be
developing as a market for sorghum-exporting countries like
the United States and Australia.
Sorghum grain produced in the Americas, Australia, and
Europe is used primarily as animal feed. It is most commonly
used in intensive production of poultry, pigs, and feedlot
cattle. (The use of sorghum grain as supplementary feed for
dairy cattle is comparatively rare.) Smaller volumes are used in
pet food. The grain is prepared in various ways for use in
different animal production systems. The simplest preparation
is to break the grain in a hammer mill or similar device and
offer as meal to animals. Due to the potential antinutritive
factors present in sorghum, this is not the most efficient use
158 THE CEREAL GRAINS | Sorghum Grain, Its Production and Uses: Overview
of the grain. More commonly sorghum is offered to poultry
and pigs in prepared complete-ration pellets and to cattle as
steam-flaked grain.
When sorghum is used in pelleted rations, it is frequently as
a cheaper substitute or complement for wheat. Due to different
gelatinization temperatures, a greater degree of heat energy
may be required to successfully bind a pellet-containing sor-
ghum. Many feed producers will add artificially produced
digestive enzymes to improve the digestibility of both the
protein and starch of the sorghum. Enzymes used in this way
include phytase, xylanase, and proteases thought to help break
down kafirins more efficiently.
Future Prospects for Sorghum
Over decades, there has been the trend in many sorghum-
producing countries for eating habits to move away from
sorghum-based foods as living standards have improved. On
the other hand, these changes may be progressively comple-
mented by increased use of sorghum for livestock feed and
alcoholic beverages (mainly sorghum beers in the same
regions) and also the likely use of sorghum as a basis for
fuel ethanol production. In addition, the specific properties
of sorghum starch may increase its use as an industrial source
of starch.
The environment favorable for sorghum production can be
expected to become greater with trends of climate change
toward warmer growth temperatures and elevated levels of
carbon dioxide in the atmosphere. Like plant breeders in gen-
eral, sorghum breeders are planning to change breeding strat-
egies. However, the expected increases in grain yield for cereals,
such as wheat (a C3 plant), are not expected to be so great for
C4 plants such as sorghum. Regarding climate change, the
greatest concern of sorghum breeders is the probable more
frequent occurrence of extreme heat events that have the capac-
ity to reduce seed set dramatically.
Exercises for Revision
List the major regions of sorghum production worldwide.
How do they differ in their utilization?
What climatic and soil conditions suit these places to sor-
ghum production? Discuss rainfall, temperatures, day
length, and latitude with respect to stages of plant growth.
How have plant breeding and selection changed the origi-
nal growth conditions for sorghum?
What climate conditions encourage or deter the diseases
and pests of sorghum?
What agronomic management can be used to reduce the
risks of damage by diseases and pests?
Describe the range of foods for which sorghum is used.
How can sorghums resistance to digestion be mitigated for
humans and animals?
Exercises for Readers to Explore the Topic Further
The protein complex of the sorghum grain is made up of
many proteins. What are their names and possible
functions?
Sorghum varieties differ in grain color. Find out the chem-
ical reasons and the possible advantages and disadvantages
of these differences.
Sorghum, like sugarcane and maize, has a C4 photosyn-
thetic pathway. It is thus distinct from C3 cereals such as
wheat and barley. What is the significance of this distinc-
tion for sorghum production now and into the future?
See also: Agronomy of Grain Growing: Sorghum: Production and
Improvement Practices; Grain Harvest, Storage and Transport:
Sorghum: Harvest, Storage, and Transport; Non-wheat Foods:
Sorghum: Utilization; Proteins: The Protein Chemistry of Cereal
Grains; The Basics: Taxonomic Classification of Grain Species; The
Cereal Grains: An Overview of the Family of Cereal Grains Prominent
in World Agriculture; Cereals: Domestication of the Cereal Grains.
Further Reading
Anonymous (2012) The U.S. Sorghum Industry. Washington, DC: US Department of
Agriculture.
Dendy DAD (ed.) (1994) Sorghum and Millets: Chemistry and Technology St Paul,
MN: AACC International.
Doggett H (1988) Sorghum, 2nd edn. London: Longman Scientific & Technical.
Pereira TD (2012) Sorghum: Cultivation, Varieties & Uses. New York: Nova Science
Publishers Inc.
Ramirez JA and Manuel Vazquez M (2012) Sorghum: Food & Energy Source. New York:
Nova Science Publishers Inc.
Smith CW and Frederiksen RA (2000) Sorghum: Origin, History, Technology, and
Production. New York: Wiley.
Relevant Websites
faostat.fao.org FAOSTAT.
www.icrisat.org ICRISAT.
 Rye Grain: Its Genetics, Production, and Utilization
HD Sapirstein and W Bushuk, University of Manitoba, Winnipeg, MB, Canada
2016 Elsevier Ltd. All rights reserved.
This article is a revision of the previous edition article by W. Bushuk, volume 3, pp. 8591, 2004, Elsevier Ltd.
Topic Highlights
The origins and history of rye production and uses are
discussed.
Rye production worldwide is falling.
International trade in rye is about 13% of world
production.
Starch (stored in discreet granules) is the major component
of rye endosperm.
Secalin is the major protein of the rye grain, but rye is not as
suitable for bread making as wheat.
The rye grain contains more fiber than wheat and other
cereal grains.
Rye contains some compounds with antinutritional prop-
erties for monogastric animals.
Rye endosperm has a soft texture, releasing flour readily in
milling.
Breads may be made from 100% rye flour but up to 90%
wheat flour may be added.
Baked goods include rye bread, rye rolls, crisp bread, and
pumpernickel.
Learning Objectives
To recognize the place of rye within the range of cereal
grains and to understand ryes genetics, morphology,
composition, and utilization.
Introduction
Cereal rye (Secale cereale L.) is an annual grass plant that is
typically fall-sown and cultivated worldwide. Historically, rye
was a major food crop in eastern European countries in the
Middle Ages due to its ample winter hardiness and lower soil
moisture requirements compared to wheat. Rye grain is used
mainly for feed and food and also for ethanol production.
Cereal rye is also used as a winter cover crop because it grows
quickly in the fall to produce ground cover that protects soil
from erosion. Cultivated area and production worldwide have
declined considerably in the past half century and currently
stand at about 6 Mha and 15 Mt, respectively (200812 aver-
age). Rye is second after wheat for production of bread, and the
grain contains the highest levels of fiber among all cereal crops.
Rye is also used as a grazing or forage crop and for silage. The
grain is also important in the production of mixed feeds for
livestock and as feedstock for the distillation of rye whisky.
Unprocessed straw is used in livestock feeds as low-digestibility
roughage and as bedding in animal husbandry. Rye straw can
also be used as a building material, for example, roof thatch
and particle board, and for other applications.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00017-6
Center of Origin and History
The primary center of origin of the rye plant is considered to be
in the Anatolian Plateau of the Middle East (Asia Minor),
encompassing the eastern region of Turkey and neighboring
portions of what is now Georgia, Armenia, Azerbaijan, Iran,
Iraq, and Syria. This northern region of the Fertile Crescent is
also the center of origin of wheat, barley, oats, and lentils.
There is no evidence of rye cultivation in ancient Egypt. Cereal
rye is believed to have diverged genetically from related Secale
species about 10 000 years ago. Rye grain is mentioned in early
writings of northern Europe suggesting this region as an impor-
tant area of historic cultivation. Cultivation of rye migrated
from its center of origin in the Middle East to northern Europe
in the first millennium BC. One possible route of migration
was from Asia Minor to western Russia and, from there, west-
ward to Poland and Germany where it is highly adaptable and
popular with farmers. Another possible route of rye plant
migration was from Turkey by way of the Balkan Peninsula to
north-central Europe. Rye was brought to North America and
South America by European settlers during the sixteenth and
seventeenth centuries. At about the same time, its cultivation
spread eastward in Europe to Siberia. During the nineteenth
and twentieth centuries, its cultivation began in Argentina,
southern Brazil, Uruguay, Australia, and South Africa. Today,
rye cultivation like that of wheat is worldwide, but the largest
production is in central and eastern Europe (Table 1).
Botanical Classification
Rye belongs to the grass family Poaceae (also called Grami-
neae) and the genus Secale. The most common cultivated
species is S. cereale, which is presumed to have evolved from
the wild perennial grass of the species S. montanum. Rye, along
with evolutionarily related cereal grains barley and wheat, is
botanically classified in the same tribe of Poaceae, that is, the
Triticeae. Rye and wheat are placed in a separate botanical
subtribe, called Triticinae, on the basis of the morphology of
their inflorescences or spikes in which the grain forms after
flowering. DNA studies have indicated that barley originated
from a common ancestor before the divergence of wheat and
rye. Cereal rye should not be confused with ryegrass that
belongs to a different genus (Lolium) of the Poaceae family
and is typically used for forage and production of turf.
Worldwide Production and Trade
Worldwide production of rye grain has declined significantly
over the past decade (Table 2), a trend that has continued for
the past half century reflecting a declining area of harvested
159
160 THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization

Table 1 Rye production worldwide (in thousands of tonnes)

Country 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012

Germany 2277 3830 2794 2644 2698 3744 4329 2903 2521 3893
Poland 3172 4281 3404 2622 3126 3449 3713 2852 2601 2875
Russian Federation 4152 2872 3628 2959 3909 4505 4333 1636 2971 2132
Belarus 1151 1397 1155 1072 1305 1492 1227 735 801 1082
Ukraine 624 1593 1054 584 563 1051 954 465 579 677
China 709 600 554 350 700 645 630 570 680 659
Denmark 169 146 132 130 135 152 238 255 294 384
Turkey 240 270 270 271 241 247 343 366 366 370
Canada 327 398 330 383 252 316 281 232 195 340
World total 14 610 17 695 15 165 12 650 15 174 18 127 18 294 11 963 13 029 14 562

FAOSTAT data, 2014. Food and Agricultural Organization of the United Nations. Accessed via the web site www.fao.org.

Table 2 Worldwide grain production (in millions of tonnes)

Grain 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012

Maize 615.5 604.9 645.2 729.0 713.7 706.8 789.9 830.3 820.0 851.2 888.0 872.1
Rice 597.5 569.1 585.1 605.8 632.2 639.1 654.8 686.2 687.0 703.2 725.0 719.7
Wheat 589.8 574.7 560.1 632.1 626.7 602.3 612.9 683.0 687.5 649.5 699.5 670.9
Soybeans 178.2 181.7 190.7 205.5 214.6 222.0 219.7 231.3 223.4 265.2 262.4 241.8
Barley 144.0 136.7 142.6 153.9 138.7 139.5 134.2 154.8 151.8 123.7 133.0 132.9
Canola 35.9 34.4 36.8 46.5 50.0 48.0 51.4 57.9 62.6 60.1 62.7 65.1
Sorghum 59.8 53.5 59.0 58.1 59.6 57.7 62.7 66.5 56.8 60.0 58.4 57.0
Millet 29.0 23.9 34.8 29.6 31.0 31.9 33.7 34.4 26.1 32.6 27.2 29.9
Rye 23.3 20.9 14.6 17.7 15.2 12.7 15.2 18.1 18.3 12.0 13.0 14.6

FAOSTAT data, 2014. Food and Agricultural Organization of the United Nations. Accessed via the web site www.fao.org.

grain over this period. Rye production is significant in abso-
lute terms but ranks last of eight grain species, including
canola, grown for human food and/or animal feed (Table 2),
the position it has occupied over the past five decades. The
decline in harvested area and production of rye is mainly due
to a decrease in demand for both food and feed and,
undoubtedly, economic returns to farmers compared with
those for other grains. Rye is grown in over 30 countries
worldwide, and 9 of these (Germany, Poland, Russian Feder-
ation, Belarus, Ukraine, China, Denmark, Turkey, and
Canada) are the main producers that together represent over
85% of world rye grain production (Table 1). The four lead-
ing countries, Germany, Poland, Russian Federation and Bela-
rus, combined to produce close to 70% of world rye
production in 2012 (Table 1).
Most of the rye grain produced is used domestically. This is
reflected in the low proportion of worldwide production that is
traded internationally; that figure is approximately 13%. The
majority of rye commerce (
85%), considering both exports
and imports, is handled by 12 countries (Table 3). Based on
most recent international statistics, between 2009 and 2011,
Poland, Germany, and Canada were the leading rye exporters
that, when combined, supply 66% of the world market on
average. In the same years, the major importers of rye were
Germany, Netherlands, and the United States, which together
accounted for 52% of total imports.
Plant and Grain Morphology
The mature rye plant has a slender, tough, fibrous stem (straw),
and elongated leaves. Plant heights vary widely from about
30 cm to over 2 m. The inflorescence (spike) is long and slen-
der with stiff long awns (beards). The appearance of rye spikes
at different stages of maturity is shown in Figure 1. The pro-
truding anthers during flowering as shown in the figure tend to
be a feature of cereal grain plants, like rye, which typically
propagate by outcrossing.
Rye grain (caryopsis) is arranged in pairs alternately on a
zigzag-shaped rachis (Figure 2). The grain is covered with a
lemma, a palea, and a glume (chaff), which is normally awned.
At maturity, individual grains can be easily seen by bending the
inflorescence (Figure 3). Like wheat, rye grain is free-threshing;
the mature grain separates easily from the glume during thresh-
ing. Rye grains or kernels tend to be more slender and longer
than wheat (Figure 4) ranging in length from 4.5 to 10 mm and
in width from 1.5 to 3.5 mm. The grains are normally of grayish-
yellow color but the color can vary widely (Figure 4) depending
on rye cultivar, region of cultivation, and harvesting conditions.
As in wheat, a crease extends the full length of the ventral side of
the grain. The surface of the grain is usually shriveled and has a
rough texture. A single grain weighs about 20 mg.
Rye grains comprise three distinct morphological parts
(Figure 5). They are the starchy endosperm, bran (pericarp
 THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization 161

Table 3 Rye grain exports and imports by leading countries (in thousands of tonnes)

Exports 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011

Germany 1001 1003 954 1344 1196 1392 584 367 465 355 225
Canada 85 58 50 213 108 158 246 97 100 160 200
Denmark 136 140 44 4 4 2 3 7 16 48 105
Poland 0 2 10 74 347 64 59 79 592 403 103
Russian Federation 0 49 400 1 0 3 129 23 8 9 85
Sweden 83 44 16 2 7 5 26 13 16 54 42
Imports 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011
Germany 14 17 79 20 145 101 167 198 354 322 268
The United States 132 129 94 162 124 142 139 144 120 120 171
The Netherlands 107 173 147 246 249 121 104 153 380 211 109
Japan 339 355 406 288 282 272 215 59 65 121 88
Poland 332 13 84 16 3 18 167 54 6 5 45
Lithuania 3 45 47 13 23 18 18 3 5 24 41
Finland 37 57 21 53 80 89 50 69 60 44 38

FAOSTAT data, 2014. Food and Agricultural Organization of the United Nations. Accessed via the web site www.fao.org.

(a) (b) (c) (d)

Figure 1 Rye inflorescence (spikes) at different stages of maturity.

(a) Before flowering; (b) early-stage flowering; (c) late-stage flowering;
(d) soft dough stage close to physiological maturity. Spikes (b) and (c)
show anthers, which contain pollen, clearly protruding from the florets in
which rye kernels will develop. Photo courtesy of Dr. Jamie Larsen,
Agriculture and Agri-Food Canada, Lethbridge Research Station.

and testa or seed coat), and germ (embryo and scutellum). By

weight proportions, these three parts of the kernel comprise
86.5%, 10%, and 3.5%, respectively. When rye grains are pro-
cessed by milling, the bran and germ are typically separated
from the endosperm, which is ground into flour.
Figure 2 Rye inflorescence with two florets removed from zigzag-
shaped rachis, which is enlarged in the inset. Photo courtesy of Dr. Jamie
Larsen, Agriculture and Agri-Food Canada, Lethbridge Research Station.

proximate constituent of the grain, which includes starch and

Composition and Nutritional Properties of Rye Grain nonstarch polysaccharides. Similar to all cereal grains, starch is
the major carbohydrate component of rye. Like wheat and
The composition of rye grain (Table 4) for the most part is barley (but unlike other cereal grains), rye starch is stored in
typical of all cereal grains, with carbohydrate being the main the endosperm in mainly two types of granules, large lenticular
162 THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization
Figure 3 Rye inflorescence approaching maturity with developing
grains. Photo courtesy of Dr. Jamie Larsen, Agriculture and Agri-Food
Canada, Lethbridge Research Station.
(a) (b) (c) (d)
Figure 4 Kernels of wheat (a), blue rye (b), red rye (c), and white rye
(d). Photo courtesy of Dr. Jamie Larsen, Agriculture and Agri-Food
Canada, Lethbridge Research Station.
granules measuring about 35 mm and small spherical granules
about 10 mm in diameter. In terms of thermal properties, that
is, when starch suspensions in water are heated close to the
boiling point and then cooled, rye starch has similar properties
of gelatinization and retrogradation compared with those of
wheat and barley starch. Interestingly, cereal chemistry
research has shown that the starch fractions of rye, wheat,
and barley are essentially interchangeable; that is, they have
similar functionality in bread making.
Rye grain contains more fiber than wheat and other cereal
grains, and that fiber resides in tissue cell walls as both water-
soluble and water-insoluble polymers. That fiber component
along with its composition and related functional properties
for health or processing properties for, for example, bread
making are the principal distinguishing features of rye com-
pared with other cereal grains. The main chemical component
of rye fiber is a polysaccharide fraction that is often referred to
as arabinoxylans and sometimes pentosans owing to its
makeup of the five-carbon sugars, arabinose and xylose.
Traditionally, this fraction was called hemicellulose, a term
that is now infrequently used. Pentosans are the main distin-
guishing chemical component of rye compared with other
cereal grains. On average, the pentosan content of whole
grain rye is about 30% greater compared with that of wheat,
while for the endosperm portion of the grain from which
refined flour derives, rye contains about twice the concentra-
tion of pentosans compared with wheat (Table 4). Pentosans
contribute significantly to the functional properties of rye flour
in bread making because of their high water-binding capacity
and interaction with storage proteins, which prevents their
aggregation into gluten required for the viscoelastic properties
of bread doughs.
The mineral and vitamin content and composition of rye are
similar to those of other cereals (Table 4). Rye is considered a
good source of B vitamins thiamin, riboflavin, niacin, pantot-
henic acid, pyridoxine, and folate and an excellent source of
a-tocopherol, the vitamin E isomer with the highest biological
activity. These micronutrients are mostly removed during mill-
ing of flour, as the B vitamins are stored mainly in the aleurone
layer of bran, while the vitamin E isomers are found in the germ.
Rye has some advantages over wheat and other cereal grains
in properties related to human nutrition and health. The
higher dietary fiber content of rye in whole grain products is
beneficial for laxation and has been shown in studies to be
more effective than whole grain wheat in overall improvement
of biomarkers of bowel health. Rye is also higher in lysine
content to the extent that this amino acid is not the first
limiting amino acid in rye as it is in other cereals. Tryptophan
is considered the first limiting amino acid in rye. The stronger
flavor and gummier texture of rye-baked products are well
liked by many consumers.
Rye grain also contains compounds that have antinutri-
tional properties especially for monogastric animals where
whole grain is used in feed in high proportion. The problem-
atic constituents include soluble pentosans and b-glucans,
which increase the viscosity of gastrointestinal contents that
interferes with feed digestion, conversion efficiency, and
growth rate and also causes sticky fecal droppings especially
in poultry. These problems can be largely eliminated with the
use of appropriate enzymes (e.g., b-glucanases and xylanases),
which hydrolyze the fiber components. For swine and cattle, it
is generally recommended that rye grain comprises no more
than about 50% of the diet.
Another issue with rye is that during flowering, florets are
often infected by a parasitic fungus (Claviceps purpurea), which
replaces the developing grain with elongated dark purple to
black dense masses of mycelia, termed sclerotia, that protrude
from the spikelets at maturity. Other cereal grains and grasses
can also be infected with this fungus, but rye is most suscepti-
ble due to its open-pollination character that facilitates access
of the fungus into the flowering head. This plant disease is
called ergot, and the resulting ergot bodies contain many toxic
alkaloid compounds, most notably ergotamine and ergono-
vine, which can cause serious health problems in livestock
depending on amount consumed and continuity of feeding.
Symptoms of acute ergot poisoning can include lameness and
gangrene due to impaired blood circulation, abortions in preg-
nant animals, seizures, and death. Contaminated feeds with
sublethal doses may still lead to problems of reduced feed
intake, poor growth and performance, and loss of milk pro-
duction in lactating animals. While sporadic cases of ergotism
can occur in cattle, swine, and sheep, particularly in foraging
animals, commercially prepared feed grain rations rarely con-
tain ergot. Ergot is of little to no significance in human con-
sumption of rye products due to the low tolerance of ergot
 THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization 163

Crease
Brush

Vascular
bundle

Central
endosperm

Prismatic
endosperm

Peripheral
endosperm
Aleurone cells

0.5 mm

Scutellum
Coleoptile
Cuticle
Plumule
Epidermis
Hypodermis
Thin walled
Rcot
cells
Sheath and cap Cross cells
Cell nucleus Testa
Nucellar
Starch epidermis
0.5 mm granules Aleurone cells
Storage Tube cells
protein Peripheral
endosperm
Prismatic
Cell wall endosperm

100 m Pericarp
(a) (b)

Figure 5 Diagrammatic view of rye grain in (a) longitudinal section and (b) transverse midsection. Reproduced with permission from Simmonds, D.H.,
Campbell, W.P., 1976. Morphology and chemistry of the rye grain. In: Bushuk, W. (Ed.), Rye: Production, Chemistry, and Technology, first ed.
American Association of Cereal Chemists, St. Paul, MN, pp. 63110.

sclerotia in commercial handling and grading of rye, removal
of ergot bodies along with other foreign material by modern
cleaning equipment prior to milling of rye grain, and the
substantial inactivation of alkaloids that occurs during baking.
Genetics, Plant Breeding, and Agronomy
Essentially, all of the commercial cereal rye grown is the dip-
loid S. cereale type. Cultivated rye contains seven pairs of
chromosomes belonging to a single genome designated by
the letter R. Accordingly, diploid rye has the genome makeup
RR. This basic genomic composition of rye is similar to that
that exists in the genetically related cereal grains, wheat, and
barley, which are all botanically classified in the same tribe of
the Poaceae family, that is, the Triticeae. Like rye, barley is also
a diploid and has seven pairs of chromosomes in its genome.
Wheat is more complex as it has a polyploidy makeup com-
prising either two or three basic sets of seven pairs of
chromosomes, which provide a much larger genetic framework
for plant breeding aimed at improving adaptation and agro-
nomic performance, protein content, and other grain compo-
sition components and end-use properties. With this goal in
mind, there have been attempts to develop tetraploid rye by
doubling the number of chromosomes with the chemical
colchicine, which inhibits cell division. While tetraploidy has
been achieved in rye, results have been mixed in terms of
practical outcomes, and little progress has been made beyond
the experimental stage.
Rye, unlike most other cereal grains, is considered to be an
allogamous breeding plant; that is, it cross pollinates naturally
and extensively with other rye plants. Therefore, it is difficult to
164 THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization

Table 4 Chemical and nutrient composition of rye compared with other cereal grainsa,b

Rye Wheat Pearled or hulless barley Oat groats or oatmeal Brown rice Corn

Kernel weight, mg 1533 2545 40 2027 2529 250500

Protein, % 1015 916 10.616 15.3 8.29.6 913
Lysine, % of total protein 3.54.5 2.72.9 2.6 3.7 3.7 26
Ash, % 1.62.1 1.8 1.21.8 2.0 1.6 1.42.1
Lipid, % 23 23 1.62.4 8.7 3.2 4.38.4
Carbohydrate, %
Starch 5864 6372 5064 54.955.8 77 6178
Fiber 1517 1013 8.0 1113 3.5 9.5
Pentosan (whole grain) 6.69.6 4.36.6 1.24.7 2.0 1.42.4 5.86.7
Pentosan (endosperm) 3.64.2 1.42.5 n/a n/a n/a n/a
b-Glucan (whole grain) 2.02.6 0.60.8 3.65.9 4.46.3 0.11 0.3
b-Glucan (endosperm) 1.52.0 0.20.4 n/ac n/a n/a n/a
Free or reducing sugars 1.3 2.0 n/a n/a 0.81.5 13
Minerals, mg per 100 g
Phosphorus 359422 150540 270 502 344 290
Potassium 387520 290620 319 425 281 370
Calcium 3170 5122 27 59 25. 30
Magnesium 92130 90290 89 158 145 140
Iron 2.710 2.84.2 3.0 4.7 1.8 3.0
Copper 0.50.9 0.42.4 0.43 0.48 0.64 0.4
Manganese 2.07.5 0.526 1.3 4.5 3.4 0.5
Zinc 3.43.9 1.910 2.3 3.7 2.2 1.4
Vitamins, mg per 100 g
Thiamin 0.46 0.51.0 0.26 0.80 0.54 0.38
Riboflavin 0.180.29 0.130.31 0.09 0.14 0.08 0.14
Niacin 1.5 4.86.4 5.0 0.96 5.58 2.8
Pantothenic acid 1.0 0.770.91 0.35 1.34 1.54 0.66
Pyridoxine 0.34 0.330.47 0.32 0.24 0.68 0.53
Folate 0.052 0.056 0.019 0.06 0.34 0.03
a-Tocopherol 1.121.20 0.501.18 0.34 0.941.81 0.130.80 0.982.11
a
Dry basis, single value typical value.
b
Various sources in the cereal chemistry literature were used to compile the data in this table.
c
Value not available.

maintain genetic purity in conventional rye populations.
Owing to the close botanical and, in essence, genetic relation-
ship between wheat and rye, their cross fertilization can be
achieved relatively easily. This is the basis for the man-made
development of the ryewheat hybrid grain called triticale. Like
rye, triticale sees modest production levels worldwide but is a
significant grain for food and feed uses.
Most of cultivated rye is a fall-sown annual crop called
winter rye although spring rye is grown in some countries,
such as Canada. Rye has excellent winter hardiness and there-
fore can be grown in areas where the climate is too severe for
winter wheat. Another distinct feature of rye that contributes to
its robust agronomic performance is its drought tolerance,
which is considered to be the best among all cereal grains
and is explained by its ability to produce an extensive root
system. Rye has been used extensively as a source of disease
resistance in wheat breeding programs. The disease-resistant
genes are incorporated through a translocation of 1R chromo-
some into 1A or 1B chromosome of wheat. Many commercial
wheat varieties contain one of these two translocations.
Rye breeding efforts, whether for conventional or hybrid
rye, are typically focused on improving grain yield as the most
important objective. Other yield-related objectives include
improving resistance to drought and nutrient stress, shortness
of straw and lodging resistance, and disease resistance (snow
mold, mildew, leaf rust, Fusarium head blight, and ergot).
Improving end-use quality of rye grain for baking is another
goal of breeding efforts. The main quality characteristic that
continues to be a detriment in rye processing is its tendency to
sprout in the head under wet harvest conditions. In contrast to
wheat, rye grain developing to maturity on the plant has rela-
tively low dormancy, which leads to preharvest sprouting. The
latter results in substantially increasing levels of the starch-
hydrolyzing enzyme, a-amylase, which transfers into flour
causing very sticky doughs that are difficult to handle in com-
mercial bread making.
The most significant aspect of rye breeding to date has been
the development of commercial hybrid rye varieties with con-
siderably enhanced grain yields, up to 1520% higher on
average, compared with the best conventional open-pollinated
counterparts. Indeed, impressive grain yields of hybrid rye of
over 11 tonnes per hectare have been reported. In Germany,
where hybrid rye research originated in the 1970s, hybrid rye
varieties today comprise 70% or more of total acreage. Hybrid
breeding that is much more complex than conventional pop-
ulation breeding exploits the heterosis effect, that is, enhanced
vigor, when two carefully selected genetically uniform but
divergent inbred parents are crossed by controlled pollination
 THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization
to obtain fertile hybrid seed. Apart from the grain yield advan-
tage to producers growing hybrid rye, the breeder benefits by
being the exclusive commercial source of that seed and
through royalties.
Primary Processing of Rye Grain
Agricultural products, including grains, vary in composition
and processing quality caused by fluctuations in abiotic (e.g.,
weather) and biotic (pests) factors during the growing season.
To facilitate marketing and processing, the grain is graded,
separated, or combined into parcels of relatively uniform prop-
erties. In countries where rye grades exist, they are based on
physical characteristics of the grain and on the presence of
foreign contaminants. Test weight, a measure of bulk density
related to milling quality, is the only grading factor that is
measured objectively. All other factors are assessed subjectively
by visual inspection. Normal or straight grades usually have a
prescribed maximum moisture content, for example, 14% in
Canada and the United States. The number of grades of milling
rye varies among countries. Germany has one grade, whereas
Canada has three, and the United States has four grades.
In transforming rye grain into food for human consump-
tion, the primary processing step is milling the grain into flour,
which basically involves two multistep processing operations,
that is, a break and reduction systems. The break system breaks
open the grain using corrugated rolls to separate bran from
endosperm and involves some additional grinding of endo-
sperm into flour, while the reduction system basically involves
reducing to flour, the so-called middling particles of more or
less pure endosperm. Despite some significant differences
between the physical structure of rye and that of wheat, the
milling processes for the two grains are similar. The first step is
the removal of all undesirable contaminants. This is achieved
by a series of special cleaning machines functioning consecu-
tively. Differences in shape, size, and density are all used to
improve the efficiency of the cleaning. The final step of the
cleaning section is tempering that involves addition of water to
toughen the bran and mellow the endosperm to facilitate their
separation. Moisture levels for tempering rye are usually in the
range 14.515.5% depending on kernel texture, and this range
is about 1% lower than that used for hard spring wheat, as rye
Table 5 Rye flour grades in North America
Flour grade Extractiona
The United States
White rye 80% of straight grade rye flour,
65% extraction
Medium rye Straight grade (SG), for example,
83%
Dark rye Clears (flour removed from SG to produce white rye flour)
Rye meal (various granulations) 100%, that is, whole grain product
Canada
Light rye 7580%
Medium rye 8385%
Dark rye 9295%
a
Extraction refers to weight percentage of whole grain milled to flour.
b
Typical values.
Adapted from Bushuk, W. (Ed.), 2001. Rye: Production, Chemistry and Technology, second ed. AACC International, St. Paul, MN, 144 pp.
165
grain is softer. For the same reason, the tempering time for rye
is shorter, typically 615 h compared with 1624 h for hard
spring wheat. Because of its softer texture, rye endosperm
releases a high proportion of its weight as flour during the
break operations. Reduction is achieved by corrugated rolls
instead of the smooth rolls used in wheat milling, as smooth
reduction rolls tend to flake rye flour, which minimizes proper
flour generation. Rye milling requires a larger sifting area than
that for hard wheat types because rye flour particles tend to
clump together because of its finer particle size distribution
arising from the softer endosperm texture.
Rye mills produce a number of different grades of flour by
systematic blending of product streams to produce flours con-
taining different proportions of endosperm and bran. In North
America, usually three flour grades are distinguished based on
degree of refinement (Table 5), which can be evaluated based
on ash or mineral content, fiber content, or flour color. A
relatively refined flour such as white or light rye will contain
mostly endosperm, and as the ash (and fiber along with pro-
tein) content of endosperm is lower than that for bran, this
explains why light rye flour has lower ash (and protein) com-
pared with medium and dark rye flour (Table 5). There is no
standard of identity of North American flours compared with
the situation that exists in Europe and Germany in particular
where rye milling and baking considerations are more precise.
A modern German rye mill will produce as many as seven
different flours designated by ash content and several types of
meals processed by crushing and cutting over a range of parti-
cle size (Table 6). Regardless of where rye is milled, the main
quality parameter for flour is ash content, which has a close
relationship to flour extraction, that is, the proportion of the
whole grain that is blended into flour. Ash content in turn has
a close correspondence with flour color; that is, the higher the
ash content, the darker the flour (Table 5).
Rye for Bread Making
Bread and other baked products are the main foods produced
from rye. Many types of baked products are made including
bread from 100% rye flour or whole meal; ryewheat bread
containing at least 50% rye; wheatrye bread containing at
least 50% wheat and at least 10% rye, rye bread, rye rolls,
Ash (%) Protein (%)b
0.60.7 79
1.01.5 1013
2.53.0 1416
1.8 1015
0.70.9
1.01.2
1.31.8
166 THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization
Table 6 Rye flour grades in Germany
a
Flour grade Extraction (%) Ash content (%, dry basis)
815 7882 0.90
997 8286 0.911.10
1150 85 1.111.30
1370 90 1.311.60
1740 1.611.80
1800 (meal) 95100 >1.80
Whole-kernel flour 100
a
The grade number is based on the ash content. 815 indicates a flour with an ash
content of 0.815%.
Adapted from Bushuk, W. (Ed.), 2001. Rye: Production, Chemistry and Technology,
second ed. AACC International, St. Paul, MN, 144 pp.
crisp bread, and pumpernickel; and other dark rye breads
made by a sourdough process that involves the use of acid-
forming bacteria and sourdough yeasts. Accordingly, rye
breads vary in color, flavor, and texture in keeping with rye
flour grade and the blended proportion of wheat flour and
other ingredients that may be used in the dough formula
besides the obligatory water, yeast, and salt. Those other ingre-
dients include shortening, sugar, molasses, caramel color, but-
termilk, lactic and acetic acid sours, and caraway seeds. The
distinct taste and aroma of rye breads, particularly of the darker
and heavier types, derive from Lactobacillus fermentation of rye
flour dough, which produces lactic and acetic acids, among
other organic acids.
The proportions of rye and wheat flour used in the bread
making formula depend on the density of the bread and inten-
sity of flavor desired by the baker. A higher proportion of rye
flour will produce bread of lower volume and more intense
flavor. The reason for this is mainly attributable to the different
composition and balance of rye endosperm proteins compared
with those found in wheat, particularly in the polymeric pro-
tein fraction, which is called glutelin in rye. Unlike wheat
proteins, rye proteins cannot form viscoelastic gluten when
the flour is mixed with water to produce dough. It may also
be the case that the high pentosan content of rye flour, and the
propensity of pentosans to bind water, could be interfering
with the normal gluten-forming process as occurs in wheat
flour doughs. Regardless of the cause, rye flour doughs are
completely inelastic, tend to be sticky, and cannot be stretched
and formed like wheat flour doughs. This makes the machin-
ability and processing of rye doughs more challenging com-
pared with those of wheat. Also, rye doughs are capable of very
little carbon dioxide gas retention due to the leavening action
of yeast and bacterial fermentation in the case of sour doughs,
resulting in low-volume bread with a coarse crumb. Where
gluten proteins in wheat are primarily responsible for gas
retention and loaf volume, in rye doughs, it is the pentosan
component and the viscosity this fraction contributes when
hydrated that is likely the principal mechanism of gas reten-
tion. For all these reasons, unlike the situation in wheat where
protein content is valued as one of the most important quality
factors, the protein content of rye flour is largely irrelevant for
bread making. Hence, when rye breads of lighter texture and
color are sought by consumers, there is a need to include an
appropriate measure of wheat flour and therefore gluten-
forming proteins in the formulation.
General Uses of Rye
Worldwide, over 50% of rye grain is used for incorporation
into livestock feed. The key factors for this application are price
relative to other feed grains such as corn and barley, energy per
unit weight, and absence of toxicants. As noted in the preced-
ing text, the principal potential toxicant stems from ergot
infection. Ergot bodies must be removed or minimized by
cleaning. For rye to be used successfully as a feed grain, con-
sideration needs to be given to its relatively high level of
pentosan content. Monogastrics and younger animals in
general tolerate this fiber component less so than ruminant
animals, and generally, no more than about 30% of the feed
should contain rye for this reason. Technology has been devel-
oped in the use of enzymes and other additives to mitigate the
effects of the antinutrients such as the soluble pentosans.
Rye grain imports by some countries, for example, Japan
and South Korea, are based almost entirely on the livestock-
feed industry. For some other countries, such as Finland, rye
grain is mainly used for bread making. Worldwide, small
quantity of rye is used in nonfeed and nonfood industries.
Traditionally, rye straw is used on the farm as livestock bed-
ding. In Canada and the United States, rye is fermented and
distilled to give alcohol that is used as a rye whiskey beverage.
Rye flour is used as an adhesive in the production of wallpaper,
corrugated packaging materials, and plywood. The capacity of
the rye plant to produce a large quantity of biomass on low-
fertility soil has not been exploited by the biomass energy
industry. The pentosan component of rye is used to make
furfural, as starting material for many industrial chemicals.
Rye plant has been reported as a significant component of a
sustainable system of agriculture, for example, as a cover crop,
but its use in this way remains to be exploited.
For additional information on rye, the reader is referred to
the monographs Rye: Production, Chemistry and Technology
(2001) and the recently published Rye: Genetics, Breeding and
Cultivation (2014). Other suggestions for further reading on
rye and relevant websites are provided in the succeeding text.
Exercises for Revision
Trace the origins and history of rye as a crop and as a food.
What reasons may be given for the gradual decline of rye
production worldwide? How might hybrid rye reverse that
decline?
Contrast the positive and negative aspects of using rye grain
for
human food,
animal feed.
Describe the taxonomic and genetic relationships of rye to
other cereal species.
Why is it difficult to retain genetic purity in rye?
Exercises for Readers to Explore the Topic Further
How does rye differ from other grain species with respect to
its genetics, disease risks, grain composition, and utiliza-
tion? List the advantages offered by rye. What disadvantages
 THE CEREAL GRAINS | Rye Grain: Its Genetics, Production, and Utilization
would need to be altered to promote rye? How might
breeding assist?
Investigate the advantages offered by F1 hybrid vigor for
other grain species (e.g., maize). What advantages could be
expected for hybrid rye?
Taking pumpernickel (or another rye-based food) as an
example, devise a marketing strategy to increase the food
use of rye, based on nutritional advantages, product appear-
ance, fashion, trendiness, international culture, or any
other perceived advantages.
See also: Breeding of Grains: Wheat Breeding: Exploiting and
Fixing Genetic Variation by Selection and Evaluation; Grain
Composition and Analysis: The Composition of Food Grains and
Grain-Based Products; Grain Harvest, Storage and Transport:
Contaminants of Grain; The Nature, Causes, and Control of Grain
Diseases in the Major Cereal Species; Grains Around the World:
Grain Production and Consumption: Europe; Grain Production and
Consumption: Africa; The Basics: Taxonomic Classification of Grain
Species; The Grain Crops: An Overview; The Cereal Grains: An
Overview of the Family of Cereal Grains Prominent in World
Agriculture; Triticale: Overview; Wheat-Based Foods: Breads.
167
Further Reading
Bushuk W (ed.) (2001) Rye: Production, Chemistry and Technology. St. Paul, MN:
American Association of Cereal Chemists.
Gorton, L.A. (2009). Formulating. In: (Editors: Pyler, E.J. and Gorton, L.A.) 4th edition.
Baking Science and Technology Volume II. Sosland Publishing, Kansas City, MO,
pp. 212220.
Schlegel Rolf HJ (2014) Rye: Genetics, Breeding and Cultivation. Boca Raton, FL: CRC
Press.
Simmonds DH and Campbell WP (1976) Morphology and chemistry of the rye grain.
In: Bushuk W (ed.) Rye: Production, Chemistry, and Technology, first ed.,
pp. 63110. St. Paul, MN: American Association of Cereal Chemists.
Wrigley CW (2010) Rye and triticale: characteristics and quality requirements.
In: Wrigley CW and Batey IL (eds.) Cereal Grains: Assessing and Managing
Qualitypp. 112140. Boca Raton, FL: CRC Press.
Relevant Websites
http://www.gipsa.usda.gov/publications/pub_fgis.html Grain Inspection Handbook.
Book II. Rye (2013).
http://www.grainscanada.gc.ca/oggg-gocg/ggg-gcg-eng.htm Canadian Grain
Commission Official Grain Grading Guide. Chapter 5. Rye (2014).
http://faostat3.fao.org FAOSTAT Data Base, Food and Agricultural Organization of the
United Nations (2014).
 Triticale: Overview
BJ Furman, San Jose State University, San Jose, CA, USA
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 3, pp 298303, 2004, Elsevier Ltd.
Topic Highlights
Triticale is a man-made crop derived from hybridization of
wheat and rye.
Triticale has desirable agronomic traits including high
yield, and resistance to biotic and abiotic stresses.
Most triticale is used for animal, use of triticale flour has
been limited by quality issues, although there are opportu-
nities in new product development.
Learning Objectives
To understand the origin and development of triticale
and the underlying genetics behind this new crop devel-
opment.
To understand where and why triticale is produced, and its
agronomic advantages and limitations.
To understand the basics of triticale processing and utiliza-
tion, especially in relation to protein quality.
Introduction
Triticale is the first man-made cereal grain crop species result-
ing from the hybridization of wheat (Triticum) with rye
(Secale), the name of which combines the scientific names of
the two genera involved. This synthetic amphiploid is obtained
by chromosomal doubling after artificial crossing to produce
fertile hybrids. It is a small-seeded cereal grain that is used
for both human consumption and livestock feed. As a hybrid
species, it combines many of the better qualities of both of
its parents. Triticale possesses wheats properties for food
production and ryes adaptive properties, and under certain
conditions can out-yield both parents. This promising crop
species is grown on more than 3 million hectares (Mha) world-
wide. Furthermore, triticale is an important germplasm source
for wheat improvement, providing a vehicle to transfer desir-
able rye characteristics to wheat.
Origin and Types
The Scottish scientist Alexander Stephen Wilson produced the
first triticale in 1876. Triticale was initially developed to com-
bine the positive traits of both parent types: the vigor and
winter hardiness as well as the higher protein content of rye
combined with the higher-quality gluten and baking properties
of wheat. However, initial progress was limited by the fact that
resulting hybrid progeny was sterile. In the 1930s, the discov-
ery and use of the chemical colchicine, a natural chemical
168 Encyclopedia of Food Grains, Second Edition
extracted from the autumn crocus plant to create chromosome
doubling, overcame this sterility problem. In 1938, Arne
Muntzing from Sweden applied colchicine to wheat/rye
hybrids, obtaining fertile plants. Once a fertile hybrid was
established, it became possible to utilize modern plant-
breeding methodologies. Early varieties were primitive and
had numerous agronomic disadvantages such as low grain
yield, poor seed set, shriveled grain, excessive height, low ger-
mination, and late maturity. Triticale improvement com-
menced in the 1960s to create new and better combinations
between wheat and rye, triticale and triticale, and triticale and
wheat. Most notable were breeding programs at the Interna-
tional Center for Maize and Wheat Improvement (CIMMYT) in
Mexico and the University of California at Davis (UCD) for
spring triticales and programs in Poland and the University of
Manitoba, Canada for winter varieties.
There are two main types of cultivated triticales: octoploid
types produced from the hybridization of bread wheat, Triti-
cum aestivum L., with rye, Secale cereale L., and hexaploid types
using durum wheat, T. turgidum L., followed by chromosome
doubling of the hybrid plant (Figure 1). Octoploid triticales
(2n 56) contain the A, B, and D genomes of bread wheat and
the R genome of rye, while the hexaploid triticales (2n 42)
contain the A and B genomes of durum wheat and the R
genome of rye. However, most triticale cultivars are hexaploids.
There is a third type of triticale (2n 28) produced from the
hybridization of diploid wheat, T. monococcum (2n 14) with
rye, but is not considered to be important economically.
Although triticale is a cross between wheat and rye, it is self-
pollinating (similar to wheat) rather than cross-pollinating
(like rye). Most agronomically desirable triticales that breed
true have resulted from several cycles of improvement, and are
primarily from the durumrye crosses with some common
wheat parentage occasionally involved.
In general, triticales can be divided into three groups
(Table 1):
1. Primary triticales are the initial product of the
wheat  rye hybridization followed by doubling of chro-
mosome numbers to produce the hexaploid or octoploid
types.
2. Secondary triticales are produced by intercrossing pri-
mary triticales or by crossing a primary triticale with wheat.
3. Substituted secondary triticales are hexaploid with A, B,
or D genomes of wheat substituted for one or more R
genome chromosomes of rye.
Primary triticales are often found to be fragile, poor producers,
and genetically unstable. They are used as breeding stock to
produce the more stable and agronomically favorable second-
ary and substituted secondary triticales. Secondary triticales
can be either hexaploid or octoploid and often contain com-
plete genomes of wheat and rye, whereas substituted triticales
http://dx.doi.org/10.1016/B978-0-12-394437-5.00019-X
 THE CEREAL GRAINS | Triticale: Overview 169

Bread wheat Rye Table 1 Development and examples of primary, secondary,

(Triticum aestivum) (Secale cereale ) and substituted triticales
2n = 42, AABBDD 2n = 14, RR
Primary triticalea
(female) (male)
Common wheat X Rye Octoploid triticale
Triticum aestivum L. Secale
cereale L.
AABBDD (2n 42) RR (2n 14) ! AABBDDRR
n = 28, ABDR (2n 56)
Durum wheatX Rye Hexaploid triticale
T. turgidum L. Secale
Embryo culture
cereale L.
AABB (2n 28) RR (2n 14) ! AABBRR (2n 42)
Seedling treated with Einkorn wheatX Rye Tetraploid triticale
colchicines to double T. monococcum L. Secale ! AARR (2n 28)
chromosome number cereale L.
Figure 1 Flow diagram for octoploid triticale development showing Secondary triticaleb
chromosome number and genome identifications. The hexaploid Triticale X Triticale
type (2n 42) is produced similarly, though using durum wheat AABBDDRR AABBDD ! AABBDDRR
(T. turgidum, 2n 28, AABB) as the female parent. (Adapted with AABBDDRR AABBRR ! AABBDDRR or
permission from Qualset CO (2002). Triticale. In: McGraw Hill Encyclopedia AABBRR
of Science and Technology, 9th edn. The McGraw-Hill Companies, Inc.) AABBRR AABBRR ! AABBRR
Triticale X Wheat
AABBDDRR or AABBDD
AABBRR
never have complete rye genomes (Table 1). One advantage of AABBDDRR or AABB
the secondary hexaploid triticale is increased genomic diver- AABBRR
sity, resulting from the insertion of portions of the D genome AABBDDRR or AA
from the hexaploid wheats. Spike type is often used as a visual AABBRR
morphological marker to distinguish types. Octoploid triticale Substituted triticalec
spikes appear similar to wheat spikes, whereas hexaploid triti- A7A7B7B7D2D2R3R3
cales have more distinctive spike types (Figure 2), and are A6A6B6B6D6D6R3R3
classified as Beagle (for complete triticales) and Armadillo A7A7B6B6D7D7R1R1
(for substituted triticales). a
Colchicine treatment is given to hybrid plants to double chromosome number.
b
Products of these hybrids have variable chromosome constitutions.
c
Examples with 2n 42. Subscripts indicate the number of chromosomes present from
each genome.
Adaptation and Production Adapted from Qualset CO, EA Rupert, and JD Prato (1973) Triticale in California:
Review of current research and appraisal as a new cereal crop. In: Yang SP (ed.)
Triticale is grown using cultural practices similar to wheat and Proceedings of the International Triticale Symposium. Lubbock, TX: International Center
rye. However under some conditions, earlier planting can for Arid and Semi-Arid Land Studies.
result in better yields. It works well planted alone, as a com-
panion crop for establishing alfalfa and for interseeding into
established alfalfa, and as a double crop with corn and other
summer annuals. There are both spring and winter growth
habits depending on the parents used in the cross, with envi-
ronmental requirements similar to other winter and spring
sown cereal grains. Drought tolerance is the primary advantage
that spring triticales have over other spring cereal crops. Winter
triticale provides a high-yielding early maturing alternative
to spring triticale for short-season areas. The University of
Manitoba began the first intensive program in North America
in 1953, working mostly with durum wheatrye crosses. Since
then, triticale has been the subject of modern plant breeding
efforts for and has resulted in excellent gains in yield and
quality. Triticale most closely resembles its wheat parent but
exhibits more vigorous growth characteristics. As a hybrid spe-
cies, it contains many of the better traits from each parent
(Table 2). Triticale can combine the bread-making qualities
of wheat with much of ryes adaptive properties such as disease
resistances, drought tolerance, and adaptability to harsh soil
conditions. As a result, varieties have been produced with a Figure 2 Spike types representing substituted (left, Armadillo) and
wide adaptive range as well as site-specific adaptation. Triticale complete (right, Beagle) forms of triticale.
170 THE CEREAL GRAINS | Triticale: Overview

Table 2 Desirable characteristics of wheat, rye, and triticalea Table 3 World triticale production, 2002

Wheat Rye Triticale Country Growth typea Area (ha) % World

High-yield potential Many grains per ear High yield China SW 500 000 16.1
Large, filled grain High biomass High-quality straw Poland W 920 523 29.7
High harvest index Low-temperature growth High feed value Germany W 560 466 18.1
Tillering efficiency Winter hardiness Disease resistant France SW 269 000 8.7
Short straw Drought tolerance Stress tolerant Australia S 264 000 8.5
Sprouting resistance Disease resistance Winter hardiness Hungary W 132 000 4.3
High-energy grain Grain high in lysine High lysine content Belarus W 94 200 3.0
Czech Republic W 53 093 1.7
a
Modified from Semundo Limited (1994) Triticale The Hybrid Evolution. Cambridge: Canada SW 47 282 1.5
Semundo Limited. Denmark W 37 657 1.2
Austria W 37 621 1.2
Sweden W 30 740 1.0
Spain S 29 900 1.0
does well in regions where wheat performs poorly, such as cold Portugal S 25 000 0.8
and infertile soils, extremely sandy soils, soils with high levels Lithuania W 20 000 0.6
of boron, salty soils, acidic soils, manganese-deficient soils, Slovakia W 18 372 0.6
and dry soils. One particular concern, however, is the presence Latvia W 15 500 0.5
of ergot infection (caused by the fungus Claviceps purpurea) in United Kingdom W 14 000 0.5
some areas. Switzerland W 13 500 0.4
The first commercial triticale cultivars were released in Belgium W 12 000 0.4
United States SW 8979 0.3
1969. Today triticale is becoming a crop in its own right and
Estonia W 6847 0.2
is grown on over 3 Mha worldwide and in at least 27 countries
Netherlands W 4618 0.1
(Table 3). This crop contributes more than 10 Mt year1 to Luxembourg W 4000 0.1
global cereal production. Since its introduction, the area har- Norway W 1000 0.03
vested has increased over 7 times and amount harvested has Tunisia S 1000 0.03
increased over 18 times (Figure 3). Mexico S 850 0.03
Although it is grown throughout the globe, the countries Algeria S 3 0.0001
that produce the most triticale are China, Poland, and World 3 122 151 100
Germany. There is also significant production in Canada and a
S: spring type; W: winter type.
United States.
Sources: United Nations FAOSTATS, Statistics Canada, and United States Census of
Agriculture.

Usage
Use of triticale for human consumption has not yet become
widespread. Although triticale flour and products are available
commercially (namely in specialty markets such as health food
stores), this availability is limited. It comes in several forms
including whole berry, flakes, and flour. Whole triticale can be
cooked and used in a variety of dishes. Quality evaluations
have shown triticale grain inferior to wheat for milling and
baking, making large-scale commercial baking not feasible.
Triticale flour is low in gluten, and bread made from it alone
is heavy. For that reason, it is usually combined half-and-half
with wheat flour. If mixed with wheat or rye flour, triticale
flour can be used to make a number of breads and pastries. In
developing countries, triticale flour is often mixed with wheat
flour during wheat shortages. It is of course important that the
crop is not infected with ergot, as this is highly toxic to
humans.
Most triticale production is used for animal feed. It offers
better amino acid balance, lysine content, and higher protein,
particularly important for swine and poultry. However, triticale
has lower energy content than other grains, and feeding of
triticale must be supplemented with other grains. It can also
be used as forage, silage, or hay for ruminants, offering high
digestibility and out-yielding traditional crops in dry soils.
Care should also be taken to insure that the crop is not infected
with ergot.
Alternative uses of triticale include use as a cover crop to
prevent soil erosion and land reclamation. Triticale has also
been used in limited amounts as raw material in bioethanol
production. Ethanol plants pay a premium for triticale over
barley, since it has more starch and no hull, making alcohol
production more efficient.
Genetic Resources
As a synthesized species, triticale has no wild ancestors and
there are no existing landrace varieties. In addition, the actual
wheat and rye parents used in triticale synthesis are often either
unknown or no longer available. It is therefore often not
possible to resynthesize unique triticale genotypes through
hybridization. Genetic resources for the development and
enhancement of triticale include existing triticales, wheat and
rye, and the ancestral species of both wheat and rye. Itself,
triticale exists as a genetic resource for the improvement of
wheat, providing a vehicle to transfer desirable characteristics
from rye.
 THE CEREAL GRAINS | Triticale: Overview 171

20 000 000
production (Mt)
area (ha)
10 000 000

8 000 000

6 000 000

4 000 000

2 000 000

0
76
78
80
82
84
86
88
90
92
94
96
98
00
02
72
74

19
19
19
19
19
19
19
19
19
19
19
20
20
19
19
19

Figure 3 World production (Mt) and area (ha) trends since the 1970s. (Source: United Nations FAOSTAT.)

In order to insure continued improvement in triticale, it is Exercises for Revision

important to maintain a comprehensive genetic resource col-
lection. CIMMYT has established a world gene bank for triticale Draw up a comparative table on the genetics, production,
and has over 15 000 accessions. The North American triticale and utilization of wheat, rye, triticale, and barley.
genetic resource collection was evaluated at the UCD and For any selected wheat flour based product, investigate the
showed a great deal of variation in both qualitative and quan- likely effects of full or partial substitution with triticale
titative traits. The collection is now maintained at the USDA flour.
Small Grains Collection in Aberdeen, ID, and at CIMMYT in
Mexico.
Exercises for Readers to Explore the Topic Further

Future Prospects What contributions are being made to triticale improve-

Triticale production has increased tremendously since the
1970s and genetic improvements have been vast. It can only
be expected that improvements will continue, especially with
the tools provided by biotechnology. In vitro regeneration of
plants will allow for successful genetic transformation. Geno-
mic maps for wheat and rye have been completed and will
provide invaluable assistance for marker-aided selection.
Although used primarily for animal culture, it holds prom-
ise on a number of additional levels. Perhaps one of triticales
greatest potential is as a vehicle for gene exchange for wheat
improvement, extending wheats gene pool. It, however, still
holds promise to be a leading food crop in some areas of the
world. Continual improvements are being made to increase
triticales grain quality for commercial production. There is a
great deal of potential for triticale products in the specialty
markets, especially in the west where a healthier and more
varied diet is becoming increasingly popular and commercial-
ized. Triticale also has potential for increasing global food
production in developing countries. It grows in many areas
unsuitable for wheat production and can out-yield wheat in
certain areas. Already used in many of these countries to some
degree, increased production for food would likely find a
market, especially in areas where wheat shortages are preva-
lent. Triticale may also play an ecological role in the future,
both for soil reclamation and biogas usage.
ment through modern genomic information resources?
What are current markets and demand for triticale? What
are likely future trends?
See also: Breeding of Grains: Wheat Breeding: Exploiting and
Fixing Genetic Variation by Selection and Evaluation; Food Grains
and the Consumer: Consumer Trends in Grain Consumption;
Genetics of Grains: Wheat Genetics; Non-food Products from
Grains: Cereal Grains as Animal Feed; Proteins: The Protein
Chemistry of Cereal Grains; The Basics: Taxonomic Classification of
Grain Species; The Cereal Grains: Rye Grain: Its Genetics,
Production, and Utilization.
Further Reading
Darvey NL, Naeem H, and Gustafson JP (1991) Triticale: Production and utilization.
In: Kulp K and Ponte JG Jr. (eds.) Handbook of Cereal Science and Technology
New York: Marcel Dekker.
Forsberg RA (ed.) (1985) Triticale CSSA Special Publication Number 9. Madison, WI:
Crop Science Society of America.
Furman BJ, Qualset CO, Skovmand B, Heaton JH, Corke H, and Wesenberg DM (1997)
Characterization and analysis of North American triticale genetic resources. Crop
Science 37: 19511959.
Guedes-Pinto H, Darvey N, and Carnde VP (eds.) (1996) Triticale: Today and Tomorrow
Boston, MA: Kluwer Academic.
Gupta PK and Priyadarshan PM (1982) Triticale: Present status and future prospects.
Advances in Genetics 21: 255345.
172 THE CEREAL GRAINS | Triticale: Overview
International Board for Plant Genetic Resources (1985) Descriptors for rye and triticale.
Rome: IBPGR Secretariat.
Limited Semundo (1994) Triticale The Hybrid Evolution. Cambridge: Semundo Limited.
Lorenz K (1974) The history, development, and utilization of Triticale. Critical Reviews
in Food Technology 5(2): 175280.
MacIntyre R and Campbell M (eds.) (1973) Triticale Proceedings of an International
Symposium Ottawa, Canada: International Development Research Center.
Muntzing A (1979) Triticale Results and Problems. Berlin, Germany: Verlag Parl Parey.
National Research Council (1989) Triticale: A Promising Addition to the Worlds Cereal
Grains. Washington, DC: National Academy Press.
Qualset CO (2002) Triticale, McGraw Hill Encyclopedia of Science and Technology, 9th
edn. The McGraw-Hill Companies, Inc.
Qualset CO, Rupert EA, and Prato JD (1973) Triticale in California: Review of current
research and appraisal as a new cereal crop. In: Yang SP (ed.) Proceedings of the
International Triticale Symposium Lubbock, Texas: International Center for Arid and
Semi-Arid Land Studies.
Tsen CC (ed.) (1974) Triticale: First Man-Made Cereal. St. Paul, MN: American
Association of Cereal Chemists.
Yang SP (ed.) (1973) Proceedings of the International Triticale Symposium Lubbock,
TX: International Center for Arid and Semi-Arid Land Studies.
 Oats: Overview
PK Zwer, South Australian Research and Development Institute (SARDI), Urrbrae, SA, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The history of oat including speciation, domestication, and
early cultivation
Oat plant growth and morphology
Growing oat including agronomy, soil factors, and climate
Growing oat diseases and pests
Processing oat
Using oat for feed, human consumption, industrial,
nonfood, and fodder end uses
Breeding improved oat varieties
Learning Objectives
To become familiar with the development of oat as a culti-
vated crop
To gain general knowledge of the morphological parts of an
oat plant and developmental stages
To learn production constraints such as agronomy, dis-
eases, and pests
To understand basic processing of oat for human
consumption
To become familiar with the many uses of oat grain and
fodder
To know the basic knowledge of oat variety improvement,
breeding systems, and new breeding technology
Introduction
Oat has played a significant role in farming systems from
domestication to the present due to the versatile uses of the
grain and plant (Figure 1). Oat currently ranks seventh in
world production of cereals after maize, rice, wheat, barley,
sorghum, and millet. World oat production exceeded rye and
triticale.
Oat is primarily grown in cool temperate climates with

67% of world production occurring in the northern hemi-
sphere. The Russian Federation, Canada, Poland, Finland,
Australia, and Finland were ranked as the top six countries
for world oat production. Oat is also grown in the southern
hemisphere with Australia ranked first in production while
Argentina, Chile, and Brazil are also significant producers.
There are two types of oats husked oat, with hulls sur-
rounding the kernel or groat after harvest, and naked oat,
where the hull is removed when the crop is harvested. Naked
oat has the free threshing character similar to wheat. Husked
oats represent the majority of global oat production except in
China where naked oat is the most common type. Naked oat is
gaining prominence for specialist markets as improved varie-
ties are being developed.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00013-9
Oat is used for animal feed, human consumption, and
nonfood uses. Figure 2 shows the uses of oat relative to the
proportions utilized.
Oat was a traditional feed on farms for centuries and pow-
ered workhorses until the introduction of machinery powered
by fossil fuels. Between 50% and 90% of the world oat pro-
duction is used as animal feed for horses, cattle, and sheep.
Naked oat is particularly suited for poultry, pigs, racehorses,
and birds. Oat as an animal fodder is also important for
domestic and export markets. Oat can be grazed as a green
feed, made into silage, utilized as straw, or baled for hay.
Oat for human consumption is used to produce traditional,
functional, and medicinal products. Oat is differentiated from
other cereal grains by using the entire kernel after the hull is
removed for many food products. Porridge or oatmeal, hot
cereals, bread, biscuits, infant food, and muesli or granola
bars are a few examples of food products produced from oat.
Nondairy food uses have been developed resulting in oat milk,
yogurt, and ice cream. Oat has been shown to have health
benefits for lowering blood cholesterol, normalizing blood
glucose levels, and reducing the risk of colorectal cancer. Phar-
macological properties are reported in the literature and prod-
ucts are being developed for commercialization.
Nonfood and industrial oat products vary from cosmetics,
to the production of cardboard products, to the manufacture of
furfural and furan compounds used for solvents, adhesives,
filtering aids, and the construction of board material and cel-
lulose pulp.
Although oat production is declining globally, specializa-
tion for animal feed, hay, food, industrial, and pharmaceutical
products is growing and provides opportunities for adding
value to the oat crop.
This article will briefly outline the significance of oat as a
cereal crop including origin, domestication, plant morphol-
ogy, production, end uses, and variety improvement.
Classification
The genus of oat is Avena L. (Poaceae) and belongs to the tribe
Aveneae of the family Gramineae. The primary species culti-
vated is Avena sativa. However, Avena byzantina and Avena
strigosa are also grown in some regions for animal feed and
fodder.
The species described in Avena form a polyploid series vary-
ing from 1 to 3 chromosome sets with the basic chromosome
number (n) 7. Each chromosome set has genome designations
A, B, C, and D donated by a different oat species in the evolution
of the genus. The different species with varying ploidy levels
evolved over time and will be discussed in the next section.
The diploid species have one set of chromosomes and are des-
ignated as 2n 2x 14. This designation indicates the diploid is
composed of one genome with 14 chromosomes. The tetraploid
173
174 THE CEREAL GRAINS | Oats: Overview
Figure 1 Oat plants pictured at heading.
Figure 2 The main end uses of oats are shown with the largest
proportion used for animal feed, the second major use for human
consumption, and minor uses for industrial, cosmetic, and
pharmaceuticals.
species have two sets of chromosomes and are designated
2n 4x 28. This designation indicates the tetraploid has two
different genomes each with 14 chromosomes, totaling 28 chro-
mosomes. The hexaploid species include cultivated oats and
have three sets of chromosomes designated 2n 6x 42.
This designation shows that the hexaploid species are composed
of three genomes each with 14 chromosomes, totaling 42
chromosomes.
The History of Oat
Speciation
Three distinct diploid genomes A, C, and D hybridized in the
development of cultivated oat. Each genome was composed of
14 chromosomes. The specific species that hybridized to form
cultivated oat has been difficult to ascertain. Several species
have been described with the A and C genomes, but no species
have been identified with the D genome. Nonetheless, domes-
ticated oat evolved from a hexaploid wild oat species com-
posed of A, C, and D genomes with Avena sterilis, Avena fatua,
or Avena hybrida all hypothesized as the progenitors of domes-
ticated oat.
Domestication and Early Cultivation
Although it is not certain where the center of origin for oat
is located, the greatest genetic diversity encompasses the
Canary Islands, the Mediterranean, the Middle East, and the
Himalayan region.
Avena species were identified at several sites in the Near East
dating from 10 500 to 5750 BC. The Neolithic Revolution
spread from the Near East to the European continent, Great
Britain, and east to Asia. By 2000 BC, farming and trading
prospered in Europe. Wheat and barley seed moved into
these regions bringing oat and rye as weed contaminants.
Although the historical record is limited, cultivated oat was
identified in the northern regions of western Europe between
4500 and 400 BC when the region experienced cool and wet
climatic changes. Oat and rye were favored in these conditions
compared to wheat and barley. There are references about oat
for fodder, animal feed, human food, and medicinal properties
by Greek and Roman authors
AD 23AD 79.
Oat cultivation continued during the Dark Ages, and by the
Renaissance, oat was ranked fourth in importance after wheat,
barley, and rye. Oat was the dominant crop in Scotland by the
thirteenth century. Oat could flourish in areas where wheat
and barley produced marginal yield and became known as a
crop that could be produced on less productive land. Although
the primary use of oat was for animal feed, by 15001700, it
was the principal grain crop for human consumption in
Scotland, Wales, Ireland, and Britain. Oat also continued to
be an important feed for horses, cattle, and sheep. During the
potato crop failure in Ireland during 174041, oat was used in
soup to sustain the hungry.
Immigrants and explorers from Great Britain and Spain
introduced oat into North America. The English introduced
oat into Canada, New England, and eastern United States in
15001600 as an animal feed. Scottish immigrants to North
America continued to use oat for porridge and other foods. The
 THE CEREAL GRAINS | Oats: Overview 175

Spanish brought oat to the Pacific coast and the southwestern

and southeastern United States in the early 1800s to feed their
horses. The general public continued to use oat for the sick,
purchasing the product at chemists. Oat production moved
west to the Upper Mississippi River Valley and into Canada
by the 1880s. Oatmeal as a breakfast food began to flourish by
1900 in United States because local mills began milling oat for
breakfast cereal, and subsequently, the product was sold in
grocery stores instead of pharmacies or drugstores.
Governor Phillip introduced oat into New South Wales,
Australia, in 1791. The primary use of oat was green fodder
and hay for horses, dairy cattle, and pigs. Oat varieties, intro-
duced from Europe, were late-maturing in Australia, so suc-
cessful production was primarily in Tasmania and the elevated
regions of mainland Australia until the early-maturing variety
Algerian was introduced for lower rainfall regions. Human
consumption of rolled oat was associated with immigrants
from Great Britain. Leonard and George Parsons emigrated
from England to Australia in 1861 and, by the late 1880s,
began manufacturing the John Bull brand of rolled oat. In
1854, Harry Clifford Love emigrated from Dublin to Australia.
After several businesses, Harry and his son Joseph formed the
Imperial Manufacturing Company, Ltd., which began produc-
ing Uncle Tobys, a brand of rolled oat in 1893.

Plant Growth and Morphology

Oat is an annual crop with tall and short stature depending on

the presence of dwarfing alleles. Each plant produces about five
stems depending on the growing season, and each stem pro-
duces about five to six leaves on dwarf stature plants and eight
to ten leaves on tall plants (Figure 3). Each stem or culm
produces a terminal panicle where the seeds develop. Plant
height will vary with growing season and the presence or
absence of dwarfing alleles. Varieties with dwarfing alleles
will vary in height between 45 and 70 cm in Australia. Tall
varieties without dwarfing alleles will vary in height between
70 and 135 cm in Australia. Root development is fibrous and
will vary according to aboveground growth and maturity but
is
1 m deep.
Figure 3 The main morphological parts of an oat plant.
Temperature and day length influence the length of the
growing period for producing oat grain. Winter and spring
oat differ by vernalization requirements. Winter oat requires
cold temperatures to produce grain, whereas spring oat will resulting in increased height of the plant. Preanthesis panicle
produce grain without the vernalization period. Due to mild development occurs when the growing point is <1 mm. As the
winter temperatures in the Mediterranean climates of the internodes elongate, the developing panicle grows upward.
southern and northern hemispheres, spring oat can be sown Unlike wheat, barley, and rye, oat forms a panicle composed
in the winter season. of branches with the seed produced at the tip. Flowering or
The major developmental stages of plant growth are germi- anthesis occurs when pollen is shed on the feathery stigmas
nation, leaf production, tiller production, stem elongation, enclosed by the lemma and palea with outer tissues called
panicle development and emergence, anthesis, grain filling, glumes (Figure 4). This stage occurs when the panicle has
and ripening. When the seed germinates, starch reserves in fully emerged from the flag leaf in tall varieties but may occur
the seed provide energy for root and leaf development until while the head is still contained in the flag leaf for dwarf
the plant begins to photosynthesize. Leaves develop at regular varieties. Fertilization occurs within 24 h. Grain size and
intervals until panicle emergence. Tiller production begins weight increase as sugars are converted to starch. As the seed
when the seedling has three to four leaves. The number of matures, the plant begins to lose moisture and senesces.
tillers that live to produce grain is dependent on environmen- The mature oat grain consists of a groat or caryopsis tightly
tal conditions and the age of the tiller. Internodes of the plant covered by a hull or husk, previously the lemma and palea
begin to elongate when about four to seven leaves are present, (Figure 5). The hull represents
3040% of the total grain
176 THE CEREAL GRAINS | Oats: Overview
Figure 4 The plant parts that comprise an oat floret. (Reproduced with
permission from Marshall HG and Sorrells ME (eds.) (1992) Oat Science
and Technology, Agronomy number 33, p. 72. Madison, Wisconsin:
American Society of Agronomy.)
weight. It is composed of cellulose, hemicellulose, and lignin.
Compared with other cereals, the oat groat is slender and
covered with hairs or trichomes under the hull. There is a
groove on the inner surface of the groat.
The groat is composed of three major fractions, bran, endo-
sperm, and germ or embryo. Several layers of compressed
tissue and aleurone cells constitute the bran located in the
outer layers of the groat (Figure 5). The aleurone cells represent
the largest component of the bran and play a role in seed
germination. The endosperm represents from 55% to 80% of
the groat. It is composed of starch, protein, lipids, and the
major concentration of b-glucans. The endosperm provides
nutrients for the growing embryo. The embryo has three struc-
tures, the scutellum, plumule, and radicle. The scutellum is
located between the embryo and endosperm and is involved
in germination and food transfer. The plumule has two to three
leaf primordia and the radicle has two to three root primordia.
Growing Oat: Agronomy
Good management practices are necessary to realize the genetic
potential of oat varieties. Sowing reliable seed, using optimum
Figure 5 Composition of an oat kernel. (Reproduced with permission
from Youngs VL (1986) Oat lipids and lipid-related enzymes. In: Webster
FH (ed.) Oats: Chemistry and Technology. St. Paul, MN: The American
Association of Cereal Chemists.)
sowing and fertilizer rates, and timely disease and pest control
constitute good management practices.
Soil Factors
Oat tolerates a wide range of soil types. The crop will tolerate
acid soils to a pH of 4.5 and alkaline soils to a pH of 8.5, but
optimum productivity occurs with pH range from 5 to 6. Acid
soils can be associated with aluminum toxicity in oat. Oat is
not as tolerant of salt as wheat, barley, and rye, but is more
tolerant than sorghum. Australian oat varieties are more toler-
ant than barley to boron, but slightly less tolerant than wheat.
Oat is generally tolerant of high manganese levels in the soil.
Climate
The most significant climatic factors affecting oat productivity
are temperature and moisture. Oat flourishes in cool moist
environments and requires more moisture to produce a unit
of dry matter than other cereals. Oat-producing regions in
North America, Europe, and Asia are primarily located between
40 N and 60 N latitudes. Maritime climates in northern
Europe are also oat-producing regions. The length of the grow-
ing season in the northern hemisphere varies from 90 to 110
days with decreasing day length as the growing season pro-
gresses. The prime oat-growing regions in the southern hemi-
sphere occur within the latitudes 20 S and 45 S. Oat is grown
 THE CEREAL GRAINS | Oats: Overview 177

between 30 S and 40 S latitudes in Australia, between 25 S

and 45 S latitudes in New Zealand, and between 20 S and
30 S latitude in South America. The growing season in Austra-
lia and South America can vary from 150 to 180 days with
increasing day length as the growing season progresses.
Although oat tolerates cold in seedling and tillering stages,
yield loss can result once the panicle emerges. However, oat
tolerates frost better than wheat and barley. Hot dry weather
can also reduce grain yield and quality especially from anthesis
to grain filling.

Agronomy
Spring varieties are primarily sown in the southern hemisphere
during the winter months. Both spring and winter varieties are
grown in the northern hemisphere and the time of sowing will
vary with latitude. Sowing rates are dependent on a combina-
tion of climate, soil conditions, and use of the crop. Sowing rates
are higher for fodder production when compared to grain pro-
duction. Seed size also varies extensively in oat. Hence, sowing
rates need to be calculated as the number of seeds m2 rather
than the weight of seeds m2 to achieve optimum plant density.
Adequate nutrition of oat is essential to achieve maximum
yields. Inputs such as nitrogen, phosphorus, potassium, and
sulfur are applied based on soil tests. Micronutrient deficien-
cies for copper, zinc, manganese, molybdenum, and iron occur
in some soils. Micronutrient toxicity for boron, aluminum, and
manganese also occurs in some soils.
Oat generally tends to be more sensitive to herbicides than Figure 6 Oat stem rust.
wheat or barley. Sensitivity is also variety-dependent.
A harvester is used to cut standing plants and threshes the
grain from the plant. Wind rowing is another option for har-
vest and requires that plants are cut and put into rows for grain
threshing at a later date.

Growing Oat: Diseases and Pests

A wide range of pests and diseases caused by fungi, bacteria,

viruses, nematodes, and insects affect all stages of plant growth,
resulting in reduced grain yield, dry-matter production, and
decreased grain quality. Control measures include genetic
resistance, chemical control, and management practices. This
brief overview summarizes the major diseases and pests in oats.

Diseases Caused by Fungi

Powdery mildew caused by Erysiphe graminis D.C. ex Marat
f. sp. avenae Marshal is an important disease in cool humid
climates such as northwest Europe. Stem rust caused by Pucci-
nia graminis Pers. f. sp. avenae Erikss. & Henn. (Figure 6) and
crown or leaf rust caused by Puccinia coronata Cda f. sp. avenae
Erikss. (Figure 7) are the two most devastating foliar diseases
in oat. Significant grain yield and quality losses result from
infection. It occurs in both the northern and southern hemi-
spheres where oat is sown. Genetic resistance is the preferred
means of control for these diseases.
Although septoria leaf blotch (caused by Stagonospora ave-
nae) is considered a minor disease compared to stem and leaf Figure 7 Oat leaf rust.
178 THE CEREAL GRAINS | Oats: Overview
Figure 8 Septoria on oat plants.
rust, the disease causes significant yield losses when it occurs.
The disease is reported in eastern Canada, the United States,
Australia, Europe, Great Britain, and Israel. Septoria is a major
production constraint in Western Australia (Figures 8 and 9).
Pyrenophora leaf blotch caused by Pyrenophora avenae Ito &
Kuribayashi apud Ito occurs worldwide with varying economic
significance (Figure 10). Pyrenophora was the third most
important disease in Germany, significant in Brazil, and com-
mon in Scandinavia. Red leather leaf caused by Spermospora
avenae Sprague & A.G. Johnson is a minor disease identified in
northwestern United States, Turkey, and Australia (Figure 11).
Diseases Caused by Bacteria
Halo (Pseudomonas syringae pv. coronafaciens) and stripe blight
(P. syringae pv. striafaciens) are collectively called bacterial
blight (Figure 12). The disease is common while cool, moist
conditions persist. Severe foliar symptoms develop on suscep-
tible varieties. Dry conditions limit the spread of bacterial
blight.
Diseases Caused by Viruses
Barley yellow dwarf virus is the most yield-limiting viral disease
of oat and is an economic threat worldwide (Figures 13 and
14). The virus is transmitted by a number of aphid species.
Figure 9 Septoria on oat leaves.
Nematodes
Cereal cyst nematode (CCN), Heterodera avenae Wollenweber,
causes a serious economic threat to oat production worldwide
(Figure 15). In regions where CCN is abundant, genetic resis-
tance reduces CCN population sizes and oat varieties with
genetic tolerance ensure maximum productivity. Genetic resis-
tance and tolerance are independently inherited.
Stem nematode, Ditylenchus dipsaci (Kuhn) Filipjev, limits
oat production in cool moist climatic conditions (Figure 16).
Winter oat sown where mild winters occur and spring oat sown
in the winter (Mediterranean climates) are affected by the
nematode. Resistance and tolerance appear to be more closely
associated for stem nematode than CCN.
Root lesion nematode, Pratylenchus neglectus, affects oat
crops in Europe, Iran, the United States, and Australia. Yield
losses up to 37% were recently demonstrated in South Austra-
lian trials where the nematode population was high. Resistance
and tolerance mechanisms are inherited similar to CCN.
Insect Pests
Russian wheat aphid (Diuraphis noxia), greenbug (Schizaphis gra-
minum), bird cherry-oat aphid (Rhopalosiphum padi), grain aphid
(Macrosiphum avenae), and rose grain aphid (Metopolophium dirho-
dum) can attack oat with varying effects on productivity.
Army worm (Phalaenidae spp.), fruit fly (Oscinella frit (L.)),
and wireworms (Agriotes spp.) are insect pests that affect oat
production in varying capacities worldwide.
 Figure 12 Bacterial blight in oat.

Figure 10 Pyrenophora leaf blotch of oat.

Figure 13 Barley yellow dwarf virus causing blasted florets and leaf
Figure 11 Red leather leaf in oat. discoloration.
180 THE CEREAL GRAINS | Oats: Overview
Figure 14 Barley yellow dwarf virus causing leaf discoloration, seed
blast, and dwarfing.
Figure 15 Cereal cyst nematode-affected plants are yellow and poorly
developed (six rows in left plot are intolerant compared with six tolerant
rows in plot behind).
Processing Oat
In order for oat to be processed, the hull must be removed from
the groat, contaminants removed, and a product produced with
quality appearance and taste. There are specifications for milling
oat that include hull to groat ratio, color, and flavor.
Milling and processing operations consist of cleaning, grad-
ing, hulling, hull and fine separation, groat separation, and
Figure 16 Symptoms of stem nematode in the left and middle plant
compared with a tolerant plant on the right.
kilning (Figure 17). Grading occurs when groats are separated
into two or three streams based on groat length or thickness.
The hulling process is the next step when groats are separated
from the hull using either impact- or stone-hulling systems.
Impact hulling is more common than stone hulling. Hulls and
fines are separated from whole groats, broken groats, and
unhulled oats. Further refinement occurs when groats are sep-
arated based on physical characters, such as groat size and
weight. Because oat groats have a high oil content, a heat
treatment is required to inactivate enzymes that cause rancidity
and bitterness in the final product. Kilning is a process that
heats the groat at a certain temperature and moisture content
to inactivate the enzymes. Kilning can occur before grading or
after groat separation (Figure 17).
Commercial processors generally can produce 100 kg of
product from 175 kg of oats. Milling efficiency varies according
to the variety and the mill-operating efficiency. Products pro-
duced include steel-cut groats, rolled oat, quick oat, baby oat,
instant oat flakes, oat flour, and oat bran.
Using Oat
Oat is a versatile crop used as grain and fodder for animal feed,
human foods, industrial products, cosmetics, and pharmaceu-
ticals. Figure 2 shows the diversity of oat products relative to
 THE CEREAL GRAINS | Oats: Overview 181

Oats corn, wheat, and barley. Lignin is the primary fiber fraction of
the hull and reduces grain digestibility in animals. Lignin
Stone milling content also varies in different varieties. Although hulls with
Impact milling Cleaning Impact milling high lignin reduce digestibility, varieties with lower-hull lignin
content can have a beneficial effect for horses, cattle, and
Kilning sheep. The hull reduces digestive problems in these animals.
The hull is a major constraint as a feed grain for poultry and
Grading Grading pigs. It reduces digestibility resulting in low protein and poor
energy. Because naked oat does not have a hull, the grain
Hulling Hulling provides a good source of energy for grower and weaner pigs,
Hull and fines Hull and fines broilers, and laying hens.
seperation seperation Overall, oat is a favorable feed for ruminants such as cattle
and sheep. Oat is also the preferred feed for horses due to the
Groat Groat
separation separation palatability, digestibility, and nutritive value of the grain.
Naked oat is also used for racehorses, due to the limited
Kilning requirement of grain intake and the need for a good source of
energy.

Broken or Large
small groats groats
Cut Cutting Flaking Grinding
groats
Milling Flaking Premium Oat
oat flakes flour
Oat bran
Oat flour
Fast cooking
oats
Figure 17 The milling process of an oat kernel.
the proportion used. Animal feed is the largest use of oat; cereal
products and other human food the second most common
use. Although pharmaceuticals represent the smallest group
of products, they have the greatest potential for future growth
and value.
Oat for Feed Grain
Oat has been a traditional feed grain for centuries. Recent
advances in corn, wheat, barley, soybean, and canola as feed
grains have resulted in a worldwide decline of oat production
for feed. Despite the decline, the primary use of oats remains as
a feed grain. Prior to mechanization, oat was the primary feed
for horses that powered farm equipment. Oat is a suitable feed
for dairy and beef cattle, sheep, and horses. Although not as
prevalent, oat can also be used for poultry, pigs, cats, dogs,
birds, rabbits, bison, deer, and fish. In recent years, naked oat is
being developed as a feed grain with improved nutritional
value for markets such as weaner and grower pigs, poultry,
racehorses, and birds. The following discussion centers on the
traditional oat grain possessing a hull.
The nutrient value for animal feed is based on the propor-
tion of groat to hull. The ratio varies with variety and environ-
ment. Oat groats have a higher oil or lipid content than other
cereals, varying between 3% and 11%. The oil is composed
primarily of unsaturated fatty acids, which can alter the fatty
acid composition of the animal fat. Protein content in oat
groats varies from 9% to 15% with higher lysine content than
Oat Grain for Human Consumption
Products produced from oat are generally made with the whole
grain after the hull is removed, whereas other cereal products
are produced from grain with the germ and bran removed.
Whole oat has the highest protein and a favorable ratio of
unsaturated to saturated fatty acids compared with other
cereals. Whole oat also provides vitamins, minerals, and anti-
oxidants. The heat treatment used to stop rancidity also
enhances the unique sensory characters of oat.
The unique composition of whole oat results in healthy
products. b-Glucan content in the groat varies between 1.8%
and 7.5% on a dry-matter basis. The high level of water-soluble
dietary fiber of which b-glucans are the main component helps
to reduce high blood cholesterol and normalize blood glucose
levels. The fiber also reduces the risk of colorectal cancer. The
groat also contains vitamins and antioxidants. Lactose-
intolerant individuals can now purchase nondairy oat
products.
The main product produced from oat is porridge or oat-
meal. Other food products processed from oat are cold cereals,
infant foods, muesli or granola bars, breads, biscuits or cook-
ies, thickeners, and specialty flour. New processes were devel-
oped recently to manufacture nondairy products such as milk,
ice cream, and yogurt using oat. Antioxidants in oat flour can
be used to stabilize some milk and meat products sensitive to
fat oxidation during storage. Oat gum primarily composed of
b-glucans is used to stabilize ice cream. Oat proteins have been
used in many food products including heat-resistant chocolate,
because of viscosity and emulsification properties.
Oat Grain for Industrial and Nonfood Uses
There are many potential uses of oat grain fractions and hulls
in products such as adhesives, cosmetic products, pharmaceu-
ticals, and oil-spill cleanup. Some of the uses are proposed,
whereas the technology for other uses has been developed and
implemented.
The unique chemical composition of the oat groat is
responsible for oat starch gels having more elastic, adhesive,
and translucent properties than wheat or maize starches. Oat
starch could replace products produced from wheat or maize
182 THE CEREAL GRAINS | Oats: Overview
with the waxy starches the exception. Products currently pro-
duced from starch include brown paper and cardboard
products and coating agents for tablets in the pharmaceutical
industry.
Oat has been used in cosmetic products for some time. The
starch characteristics make oat a useful ingredient in bath
products to relieve itching and nappy rash. Other products
produced from oat include cleansers to replace soap in facial
masks, soap for cleansing dry and oily skin, shampoo, lotions,
bath additives, and skin care.
Pharmaceutical products have great potential, but are not
currently tapped. The health benefits of oat fractions were
discussed in the preceding section.
The hull can also be used to produce industrial products.
Oat hulls are used to produce furfural and furan compounds.
These compounds are used in the manufacturing process of
crude petroleum; nylons; formaldehyde furfural resins; sol-
vents for dyes, resins, lacquers, paints, and varnish; elastomers
and thermoplastics; phenolic resin glues and plywood adhe-
sives; construction board material; and cellulose pulp. A very
different aspect of hull utilization is the cariostatic properties of
tooth protection.
Oat for Fodder
Fodder production is also an important component of animal
feed shown in Figure 2. Several types of oat fodder, such as
hay, silage, grazing, and straw, are produced as an essential
component of animal production systems worldwide.
Oat hay is produced for dairy and beef cattle, horses, and
sheep. Hay not only is an important commodity domestically
but also is traded internationally. A significant market is the
Asian dairy and beef cattle industry. The oat hay crop is cut at
the watery-ripe growth stage (Zadoks 71) for optimum hay
quality. Hay quality characters that may be associated with
palatability are digestibility, water-soluble carbohydrates, acid
detergent fiber, neutral detergent fiber, and stem diameter.
Sensory quality characters are also important and include hay
color and aroma.
Cattle and sheep graze oat during the vegetative stage when
the plant height is from 20 to 40 cm. Animals are removed
from the oat crop before the growing point of the plants is
damaged, so the crop produces grain. Grazing is important in
countries worldwide including New Zealand, Australia, South
America, northern Africa, Nepal, and the United States.
Silage production using oat is used in the dairy and beef
cattle industry worldwide. Oat is cut in the early heading stage,
chopped, and ensiled.
Breeding-Improved Oat Varieties
History
Early oat improvement occurred when farmers selected the
best-performing plants in their paddocks. The first attempt to
introduce genetic variation by crossing different oat genotypes
was reported in England and accomplished by Shirreff in the
1860s. The first variety developed from hybridizing oat, Abun-
dance, was released from Gartons Ltd., Warrington, England.
Breeding Priorities
Breeding priorities determine selection criteria that eventually
result with the release of oat varieties with improved traits.
Understanding production and processing constraints of oat
is the initial step in identifying breeding priorities for improv-
ing varieties. These priorities will differ or change in emphasis
depending on the growing environment and end use of the oat
plant or grain. Breeding priorities are based on gross margin for
growers and processors, resulting in a superior product for the
consumer and processor. Breeding priorities set to achieve
long-term goals are based on developing varieties that offer
new products or increased value for both growers and industry.
The long-term breeding priorities are particularly important for
oat due to the global decline of production.
Although there are many different growing environments
and end uses globally, the primary breeding priorities deal with
productivity, disease resistance, agronomy, and quality.
Increased grain and hay yields are the primary aims of
increased productivity. Improved disease resistance increases
yield and enhances quality.
The significance of oat diseases varies with region and con-
tinent and is prioritized by breeding programs. Diseases such
as stem and leaf or crown rust are two global diseases that cause
significant losses to the oat industry. Barley yellow dwarf virus,
smut, and septoria also reduce yield and quality in certain
production areas of the world. Cereal cyst, stem, and root
lesion nematodes when present in the production environ-
ment are equally as devastating, but more regionally localized
in the world.
Important agronomic characters are standing ability, matu-
rity of the plant, and the ability of the plant to resist shattering
or loss of seed at maturity. Winter hardiness is also an impor-
tant agronomic trait of the varieties that require a cold period
to produce seed.
Grain quality characters that are essential for the industry
are high groat to hull ratio and groat color. Weather and
disease can cause groat discoloration. Other grain quality char-
acters of importance for variety development are protein, oil,
b-glucan content, hectoliter weight, and screening percentage.
The most important aspect of fodder quality is palatability. The
traits that comprise palatability are being researched but are
most likely a combination of digestibility, water-soluble car-
bohydrates, neutral detergent, and acid detergent fibers, color,
and smell.
Breeding Methods
Breeding systems should be flexible and easily modified for
changes in breeding priorities. A combination of breeding
methods is combined in a program to produce improved oat
varieties. Classical breeding methods used for decades are ped-
igree, bulk, recurrent selection, backcross, and single-seed
descent. More detailed information about the breeding
methods is available in Plant Breeding Methodologies by Neal
Jensen.
Doubled haploids (DH) are true breeding homozygous lines
produced in one generation. An economically viable method for
DH production in oat is currently not available, but new devel-
opments using isolated microspore culture hold promise for

future implementation. DH technology is used commercially
for wheat, barley, and canola breeding.
Molecular marker technologies for oat have only recently
started compared with those for wheat and barley. Molecular
markers are a selection tool using DNA to identify lines with
desirable combinations of traits. It provides fast and accurate
trait selection compared to traditional data collection in field
trials where the environment influences the phenotype. How-
ever, it is necessary to have an accurate phenotypic analysis to
establish the closest linkage between the trait and molecular
markers for marker development.
Germplasm Collections
Oat germplasm collections are maintained in many countries
and are a source of new genetic variation for improving oat.
Without these collections and the exchange of advanced germ-
plasm from oat-breeding programs, oat variety improvement
would not advance. Several collections of notable importance
are located in the United Kingdom, Canada, the United States,
Israel, the Russian Federation, and China.
Exercises for Revision
Describe the main end uses of oat and what proportion of
the crop is used for each.
Describe the history of oat utilization from the first docu-
mentation until the 1700s.
Name the morphological parts of an oat plant and oat floret.
Discuss the composition of an oat kernel.
Name three fungal diseases of oat and their causal pathogens.
Name virus and nematode diseases and their pathogens.
Discuss breeding methods including new technologies.
Exercises for Readers to Explore the Topic Further
What are the healthy components of the oat grain?
What is the current research status for molecular markers
and doubled haploids in oat?
What is the current research on the healthy aspects of oat
for animal feed?
What is happening with drought tolerance research in oat?
See also: Agronomy of Grain Growing: Plants: Diseases and
Pests; Grain Harvest, Storage and Transport: Cereals: Grain
Defects; The Nature, Causes, and Control of Grain Diseases in the
Major Cereal Species; Non-food Products from Grains: Cereal
Grains as Animal Feed; Proteins: The Protein Chemistry of Cereal
Grains; The Basics: Taxonomic Classification of Grain Species; The
Cereal Grains: An Overview of the Family of Cereal Grains Prominent
in World Agriculture; Cereals: Domestication of the Cereal Grains;
Wheat-Based Foods: Cereals: Overview of Uses: Accent on Wheat
Grain.
THE CEREAL GRAINS | Oats: Overview 183
Further Reading
Baum BR (1977) Oats: Wild and Cultivated. A Monograph of the Genus Avena L.
(Poaceae). Ottawa, ON: Department of Agricultural Supply and Services Canada.
Bonnett OT (1961) Morphology and development. In: Coffman FA (ed.) Oats and Oat
Improvement. Madison, WI: American Society of Agronomy.
Bowden CW (1987) Oats. In: Lazenby A and Matheson EM (eds.) Australian Field
Crops: Wheat and Other Temperate Cereals. North Ryde, NSW: Angus and
Robertson.
Clifford BC (1995) Diseases, pests, and disorders of oats. In: Welch RW (ed.) The Oat
Crop: Production and Utilization. London, UK: Chapman and Hall.
Cuddeford D (1995) Oats for animal feed. In: Welch RW (ed.) The Oat Crop: Production
and Utilization. London, UK: Chapman and Hall.
Ferrie AMR, Irmen KI, Beattie AD, and Rossnagel BG (2013) Isolated microspore
culture of oat (Avena sativa L.) for the production of doubled haploids: Effect of
pre-culture conditions. Plant Cell, Tissue and Organ Culture 116: 8996.
Forsberg RA and Reeves DL (1995) Agronomy of oats. In: Welch RW (ed.) The Oat
Crop: Production and Utilization. London, UK: Chapman and Hall.
Ganssmann W and Vorwerk K (1995) Oat milling, processing, and storage.
In: Welch RW (ed.) The Oat Crop: Production and Utilization. London, UK: Chapman
and Hall.
Hoffman LA (1995) World production and use of oats. In: Welch RW (ed.) The Oat Crop:
Production and Utilization. London, UK: Chapman and Hall.
Jensen NF (1988) Plant Breeding Methodology. New York, USA: John Wiley and Sons.
Leggett JM (1992) Classification and speciation in Avena. In: Marshall HG and
Sorrells ME (eds.) Oat Science and Technology. Madison, WI: American Society of
Agronomy.
Moore-Coyler RJ (1995) Oats and oat production in history and prehistory.
In: Welch RW (ed.) The Oat Crop: Production and Utilization. London, UK: Chapman
and Hall.
Murphy JP and Hoffman LA (1992) Origin, history, and production of oat.
In: Marshall HG and Sorrells ME (eds.) Oat Science and Technology. Madison, WI:
American Society of Agronomy.
Pomeranz Y (1995) Industrial uses of oats. In: Welch RW (ed.) The Oat Crop:
Production and Utilization. London, UK: Chapman and Hall.
Ranhotra GS and Gelroth JA (1995) Food uses of oats. In: Welch RW (ed.) The Oat
Crop: Production and Utilization. London, UK: Chapman and Hall.
Schrickel DJ, Burrows VD, and Ingemansen JA (1992) Oat harvesting, storing, and
feeding. In: Marshall HG and Sorrells ME (eds.) Oat Science and Technology.
Madison, WI: American Society of Agronomy.
Sidhu PK and Davies PA (2009) Regeneration of fertile green plants from oat isolated
microspore culture. Plant Cell Reports 28: 571577.
Webster FH (1986) Oat utilization: Past, present, and future. In: Webster FH (ed.) Oats
Chemistry and Technology. St. Paul, MN: American Society of Cereal Chemists.
White EM (1995) Structure and development of oats. In: Welch RW (ed.) The Oat Crop:
Production and Utilization. London, UK: Chapman and Hall.
Youngs VL (1986) Oat lipids and lipid-related enzymes. In: Webster FH (ed.) Oats
Chemistry and Technology. St. Paul, MN: American Society of Cereal Chemists.
Zadoks JC, Chang TT, and Konzak CF (1974) A decimal code for the growth stages of
cereals. Weed Research 14: 415421.
Relevant Websites
www.eatmoreoats.com This website gives information about health, recipes, and
products.
www.grdc.com.au This website gives research information in Australia and
Worldwide.
www.quakeroats.com.au This website gives information about oat products.
www.rirdc.com.au This website gives information about fodder research for oat hay
varieties.
www.taste.com.au This website gives information about oat recipes.
www.uncletobys.com.au This website gives information about oat product, health and
nutrition, and oat recipes.
www.whfoods.com This website gives information about oat nutrients, health benefits,
history, and cooking.
 Coix: Overview
H Corke, University of Hong Kong, Hong Kong, Pokfulam, China; Hubei University of Technology, Wuhan, China
Y Huang and JS Li, China Agricultural University, Beijing, China
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Native to Asia, coix is a close relative of maize and produces
a grain widely used in Chinese medicine.
The storage protein of coix largely comprises a prolamin
fraction termed coixin.
Various components of the coix grain, particularly coixol,
coixin, and coixenolide, are of medicinal and nutritional
importance.
Learning Objectives
To understand the origin and taxonomy of coix and its
relationship to maize.
To understand the uses of coix, both as food and in tradi-
tional medicine.
To understand the basics of the chemical composition of
coix that contribute to its medicinal properties.
Introduction
Coix lacryma-jobi L., commonly called Adlay, Jobs tears, adley,
adley millet, or coix millet, is an annual crop. Botanically, coix
belongs to the tribe Maydeae (Tripsaceae) of the family Poa-
ceae, and it is classified as a close relative to maize. This annual
grass is native to India, Burma, China, and Malaysia and has
been grown extensively in Southeast Asia for several thousand
years. The plant is typically
1 m high with knobbly bamboo-
like stems, and the new tillers arise from the base of stems. The
glossy deep green leaves are up to 5 cm wide with slightly wavy
edges. The flowering and fruiting spikelets are insignificant; the
grain is pear-shaped and
5 mm in diameter and has a hard,
shiny dark brown to grayblack hull. The grain is used in soups
and beverages and is popular in traditional Chinese medicine.
It is well known to have pharmacological properties. The
ancient Chinese medical book Pen-tsao kang mu described it
as an effective remedy for a number of maladies and particularly
beneficial to the digestive system. Coix has been considered as a
good potential germplasm resource for maize improvement
because it is tolerant to lateritic soils and low pH, less prone
to attacks by viruses, and less susceptible to waterlogging and
has a low variation in photoperiod. Accurate world production
figures are impossible to ascertain, but coix has a value beyond
that of a typical commodity grain for food use.
Botany
Habit
Coix is a light-sensitive and short-day-length plant and prefers
wet soils and cool weather. Although it is an annual crop, the
184 Encyclopedia of Food Grains, Second Edition
plant is perennial in essentially frost-free areas. It is reported to
tolerate annual precipitation of 6.142.9 dm, annual temper-
ature of 9.627.8  C, and soil of pH 4.58. It can be grown in a
variety of environments from cool-temperate moist to wet to
tropical very dry to wet forest life zones, forest margins, and
swamps. Although this annual grass is native to Southeast Asia,
it is now distributed throughout the tropical, subtropical, and
temperate zones.
Vegetative Organization
The coix plant is 70400 cm high. Its stems are erect or strag-
gling with branches and prop roots from the lower nodes. The
internodes of the culms are solid. Leaves are not basally aggre-
gated and nonauriculate, the leaf blades are lanceolate and
broad and
3070 mm wide, the shape is cordate or not
cordate but not sagittate, they are flat and persistent without
cross venation, they are rolled in bud with a ligule present, and
there is an unfringed to a fringed membrane.
Reproductive Organization
The coix plant is monoecious with all the fertile spikelets being
unisexual but without hermaphrodite florets (Figure 1). The
spikelets of sexually distinct forms (female-only or male-only)
are on the same plant, the glumes of the fertile spikelet are
several-nerved, and all are enclosed finally in a bony beadlike
involucre containing a minute fertile female flower and two
sterile flowers. Pollen-bearing male flowers are produced on a
slender stalk that extends out of the bead through a tiny pore.
Two feathery stigmata from the fertile female flower also pro-
trude from the pore ready to receive pollen from the male
flowers:
1. Female-sterile spikelets: Male spikelets grow in pairs or tri-
ads, several per disarticulating raceme. Rachilla of male
spikelets is terminated by a male floret. The male spikelets
have glumes and proximal incomplete florets or not (the
lower floret sterile or male).
2. Female-fertile spikelets: Spikelets fall with the glumes.
Rachilla is terminated by a female-fertile floret. Usually, it
is pollinated by the wind.
Seed
The shiny gray to black beads are dispersed and planted like
seeds, but they are like little shells containing flowers and
grains. The seeds are usually oval-shaped or egg-shaped, with
a milky white outer surface and a slightly sweet taste (Figure 2).
Fruit is medium sized with globular form and 612 mm long.
Close-up view of flowering coix shows the hollow, beadlike
involucres, which have a natural hole in them.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00008-5

Thrumb
Female spikelet
Flower stalk
Male spikelet
Figure 1 Inflorescence of Coix. Adapted from the images from http://
www.desert-tropicals.com/plants/Poaceae/coix_lacryma-jobi.html.
Figure 2 Photograph of flower and seed of Coix. http://www.biherb.
com/CoixSeedExtract.html.
Cytology
The standard karyotype formula is 2n 20 20m (Coix puel-
larum). But there are other karyotype formulas outside of the
standard types, such as 2n 20 18m 2sm (C. stenocarpa);
2n 20 18m 2sm and 2n 20 16m 4sm (Coix lacryma-
jobi var. lacryma-jobi); 2n 20 12m 2m (sat) 6sm (Coix
THE CEREAL GRAINS | Coix: Overview 185
Figure 3 The karyotype of Coix. Reproduced from Zhang F and Yi Z
(1994) Natural Science Journal of Changsha Normal University of Water
Resources and Electric Power 9(2): 187191.
lacryma-jobi var. monilifer); 2n 20 14m 6sm (1  2B)
(Coix lacryma-jobi var. formosana); and 2n 20 18m (2
sats) 2sm (Coix lacryma-jobi var. ma-yuen (Roman) Stapf). In
another research, the chromosome complement of 2n 10 and
2n 20 was also found in different populations of coix of
Indian origin (Figure 3).
Biochemistry
Like other members of the grass family, coix has seeds contain-
ing prolamin as the major storage protein. The coix prolamin is
known as coixin and represents more than 70% of the endo-
sperm protein. Based on differential solubility, coixins can be
separated into two factions: a-coixins and g-coixins. a-Coixins
are constituted by four size classes, while g-coixins comprise
only one molecular weight class. a-Coixins are synthesized at
earlier developmental stages than g-coixins. The deduced
amino acid sequence of a full-length cDNA clone encoding a
sulfur-rich coix prolamin predicted a polypeptide of 194 resi-
dues, which shared 64% homology with the 17 kDa b-zein.
The mature protein contains the familiar composition of pro-
lamins and an unusually high content of the sulfur-containing
amino acids methionine (11.6%) and cysteine (5.2%).
Hydropathy analysis showed that a-coixin is slightly more
hydrophobic than b-zein.
The anticomplementary polysaccharides CA-1 and CA-2
purified from the seed of Coix lacryma-jobi L. var. ma-yuen
both consist of rhamnose, arabinose, xylose, galactose, galac-
turonic acid, and glucuronic acid but in different ratios. CA-2
contains more mannose and glucose. CA-2 showed more
potent anticomplementary activity than CA-1 in low dose.
The seeds of C. lacryma-jobi L. var. ma-yuen Stapf are used as a
traditional Chinese medicine reputably possessing antitumor
activity, attributed to an acidic fraction. Infrared spectroscopy
and gasliquid chromatography showed that this acidic frac-
tion was composed of four free fatty acids: palmitic, stearic,
oleic, and linoleic acids.
A protein inhibitor of locust gut a-amylase that was purified
from coix seeds consists of two major isomers, each dimer of
two closely similar or identical subunits of Mr
26 400 Da and
associated by interchain disulfide bonds. These isomers
also have closely similar amino acid compositions. The major
isomer shows no inhibitory activity against amylases from
other sources (human saliva, porcine pancreas, Bacillus subtilis,
Aspergillus oryzae, and barley malt). This highly specific func-
tion may be relevant to protection of the grain from insect
feeding and fungal infection. The complete amino acid
sequence of the polypeptide of the major trypsin inhibitor
from coix seeds showed 64 amino acids with a high content
of cysteine. The sequence exhibits strong homology with a
186 THE CEREAL GRAINS | Coix: Overview

number of BowmanBirk inhibitors from legume seeds and Steady-state levels of DHPS mRNA are slightly reduced in the
similar proteins recently isolated from wheat and rice. endosperms and embryos of the maize lysine-rich O-2 mutants
The amylose contents of normal coix ranged from 15.9% to when compared with those in normal kernels. Also, the DHPS
25.8%, and those of waxy coix were 0.71.1%. Swelling power gene is not under the control of O-2.
of waxy coix starches varied between 28.6 and 41.0 g per g, Using random amplified polymorphic DNA (RAPD) markers,
generally higher than waxy maize. Normal coix starches 21 coix accessions were characterized for genetic variations and
had significantly higher gelatinization peak temperature relationships. The results indicated considerable variation at the
(71.171.4  C) than normal maize (71.975.5  C), similar to DNA level among the coix germplasm, and the classification by
waxy maize. Rapid Visco Analyser pasting profiles of normal RAPD data reflected the differences in geographic origins and
coix show some variation but closely match the normal maize evolution of coix. Recently, a coix genetic linkage map with
starch profile. Pasting profiles of waxy coix have more variation restriction fragment length polymorphism (RFLP) and amplified
and lower peak viscosities than waxy maize starch. Waxy coix fragment length polymorphism (AFLP) markers was constructed.
starches form very weak gels, while the gel hardness of normal The map consisted of ten linkage groups, consistent with the
coix starches was 11.431.1 g. The gelatinization temperature chromosome numbers observed cytogenetically.
of waxy coix starches varies between 63.4 and 76.4  C, and
water-binding capacity ranges from 103% to 108%. The starch
yield from coix grains averages
45%, the birefringence loss of Utilization
starch granules begins at 65  C and finishes at 75  C, and
breakdown during pasting in a Brabender Visco-Amylo-Graph Coix may be a source of ornamental beads, a staple sustenance,
is substantial. The properties of waxy coix starch are similar to and a productive fodder grass increasingly viewed as a poten-
those of waxy maize starch. Amylose content is the main factor tial energy source. It has long been consumed both as a nour-
controlling differences in starch properties of coix starches. ishing food and as an herbal medicine.
Oil yields ranged from 3.03 to 4.83 wt%, ferulate phytos-
terol esters in oils ranged from 0.109 to 0.119 wt%, free phy-
tosterol varied from 0.54 to 0.61 wt%, phytosterol fatty acyl Food
esters in the oil ranged from 0.79 to 0.98 wt%, total phytos-
terols were from 1.43 to 1.69 wt%, and g-tocopherol varied Before maize became popular in South Asia, coix was rather
between 0.030 and 0.045 wt%. Whole kernel oil yields of coix widely cultivated as a cereal in India and still occurs as a minor
are generally higher than Zea mays ssp. mays and teosinte cereal. Coix was used as food, particularly by peasants in the
accessions. Whole kernel oil yields and levels of four phytonu- Far East, very early in agricultural history. In wild varieties, the
trients (free phytosterols, fatty acyl phytosterol esters, ferulate fruit has a hard, shiny coat. After domestication, this coat
phytosterol esters, and g-tocopherol) in oils were measured became less hard and easier to cook, for example, into por-
from three coix accessions (C. lacryma-jobi L.). ridge. Coix ranks along with wheat and barley in the Near East;
beans, maize, squash, and pepper in the Americas; and rice in
Asia as one of the earliest domesticated plants. As with other
Molecular Biology cereals, there are many coix genotypes, including soft-shelled,
easily threshed types with a sweet kernel.
The seed storage proteins of coix, sorghum, and maize are Though the hard seed coat, removed by decortication,
coded by homologous genes, which are coordinately expressed makes extraction of the flour rather difficult, this is a poten-
in the endosperm in a temporal-specific fashion. Opaque-2 tially very useful grain because it has a higher protein to car-
(O-2), a bZIP protein originally isolated from maize, has bohydrate ratio than any other cereal (Table 1). Reported data
been described as a transcription activator of a-prolamin and differ somewhat. According to List and Horhammer, 100 g of
b-prolamin genes. The coding region of the coix O-2 gene is seed contains 358 cal, 11.2 g H2O, 15.4 g protein, 6.2 g fat,
interrupted by five introns and codes a polypeptide of 408
amino acids. Comparison of the deduced amino acid sequence Table 1 Typical composition of Coix seed compared with other
with two different sequences of maize O-2 protein showed that grains (on a 14% moisture basis)
the coix O-2 protein is similar to the maize O-2 isolated from a
Coix Rice Wheat Maize
maize inbred line, W22. The coix O-2 protein has the same
binding specificity and expression pattern of the maize O-2. Energy (cal) 1500 1460 1480 1510
Dihydrodipicolinate synthase (DHPS) is the main enzyme Protein (%) 14.2 7.8 9.9 8.4
of a specific branch of the aspartate pathway leading to lysine Fat 3.6 1.3 1.8 4.3
biosynthesis in higher plants. The open reading frame of the Starch (%) 67.2 76.6 74.6 70.2
DHPS-encoding DapA gene from C. lacryma-jobi is interrupted P (mg) 299 203 268
by two introns and encodes the 326-amino acid-long coix Mg (mg) 126
DHPS protein, which is 95% identical to the maize DHPS Ca (mg) 49.0 9 38 34
protein. DHPS transcripts are present in coleoptiles, embryos, Fe (mg) 2.90 4.2
Zn (mg) 1.89
endosperms, and roots but are almost undetectable in blades
Mn (mg) 1.89
of young leaves of both coix and maize. The 50 -flanking region
of DapA gene contains a TGACTC GCN4-like element located Per 100 g for fat and mineral contents.
372 bp upstream from the putative translation start codon. Source: www.fao.org, www.chinafeedbank.com.cn, and www.cmymy.51.net.

Table 2 Amino acid (aa) composition as % of crude protein
aa Arg Cys Gly Hys Ile
4.4 1.7 2.8 2.3 4.0
nd, not detected.
Source: www.fao.org.
65.3 g total carbohydrate, 0.8 g fiber, 1.9 g ash, 25 mg Ca,
435 mg P, 5.0 mg Fe, 0 mg b-carotene equivalent, 0.28 mg thia-
mine, 0.19 mg riboflavin, 4.3 mg niacin, and 0 mg ascorbic
acid. There is 5060% starch, 18.7% protein (mainly gluta-
mine, leucine, tyrosine, arginine, histidine, and lysine), and
510% oil with glycerides of myristic and palmitic acids. The
partial amino acid composition of crude protein in coix seeds
is shown in Table 2. It should be noted that despite the level of
lysine noted, it is still deficient relative to human dietary needs.
Coix seeds can be used for baking, mixed with wheat flour,
and for making porridge and beer. They are usually pounded,
threshed, and winnowed. The pounded flour is sometimes
mixed with water, similar to use of barley for the beverage
barley water. The pounded kernel is also made into a sweet
dish by frying and coating with sugar. The grains are also
utilized in soups, broths, porridge, drinks, and pastries. In
India, the Nagas, a group of tribes near the Myanmar border,
use the grain for brewing a beer called zhu or dzu. A Japanese
variety called Ma-Yuen is brewed into a tea and an alcoholic
beverage, and roasted seeds are made into a coffee-like drink.
In some cases, the hulled grain is adapted for parching or
boiling like rice; it is also dehusked and eaten like a peanut.
Beers and wines are made from the fermented grain. Continu-
ing recent interest in new product development from coix is
apparent, and coix bran may be particularly notable as a high-
potential food ingredient.
Medicine and the Rise of KanglaiteW
According to traditional Chinese medicine, coix seeds serve
several functions. They stimulate function of the spleen and
lung, remove heat, help in the drainage of pus, and induce
diuresis. Coix seeds have been widely employed as a diuretic,
stomachic, analgesic, and antispasmodic agent from ancient
times. They are also used to treat the symptoms of diarrhea and
arthritis. The fruits are anodyne, anti-inflammatory, antipy-
retic, antiseptic, antispasmodic, hypoglycemic, hypotensive,
sedative, and vermifugal. According to Hartwell, the fruits are
used in folk remedies for abdominal tumors; esophageal, gas-
trointestinal, and lung cancers; various tumors; and excres-
cences, warts, and whitlows. This folk reputation is all the
more interesting when reading that coixenolide (Figure 4)
has antitumor activity. It is also a folk remedy for abscess,
helminthiasis, anthrax, appendicitis, arthritis, beriberi, bron-
chitis, cancer, catarrh, diabetes, dysentery, dysuria, edema,
fever, headache, phthisis, pleurisy, pneumonia, rheumatism,
small pox, splenitis, etc., is an anodyne and tonic, and can be
used during puerperium. Coix seeds are reportedly effective in
treating verrucas caused by the human papillomavirus. A tea
from the boiled seeds is drunk as part of a treatment to cure
THE CEREAL GRAINS | Coix: Overview 187
Leu Lys Met Phe Thr Try Tyr Val
14.4 1.9 3.0 4.8 3.1 nd 4.2 5.6
warts. It is also used in the treatment of lung abscess, lobar
pneumonia, appendicitis, rheumatoid arthritis, beriberi, diar-
rhea, edema, and difficult urination. The action of coix seeds
against many kinds of disease can be attributed to various
components with pharmacologically different activities. Coix
roots can be also used in the treatment of menstrual disorders.
A major development in the utilization of coix has been the
advance of Kanglaite, which is coix seed oil injected intrave-
nously as a treatment for many forms of cancer. Coixenolide
and coixol (regarded as anticancer and anti-inflammatory,
respectively) are two major active fractions of coix oil (see
Figure 3). It has become a widely used cancer therapy, and a
much-hyped treatment in the West. Clinical evidence is posi-
tive to some degree, and many studies have recently been
conducted, for example, on gastric cancer, on pancreatic cancer
xenografts, and on stimulation of anticancer-related immune
responses. In fact, numerous recent studies indicate possible
anticancer and other medicinal roles of polysaccharide, protein
(antidiabetic), polyphenols, bran, and oil of coix. Some of
these uses may come to lack support from future rigorous
clinical trials. Some uses may depend on complex interactions
of coix components with other aspects of a long-term healthful
diet. But certainly, there is abundant evidence that coix may
have vast future potential as a nutraceutical source.
Fodder
The leaves and stems provide a useful fodder for cattle and
buffalo. The grass is very suitable for growing in waterlogged
areas. Its average yield is 29 tonnes ha1. The grass can also be
turned into silage.
Genetic Resources
Coix is reported to tolerate lateritic soils, low pH, slope,
viruses, and waterlogging and to be photoperiod-insensitive.
It is a potential gene resource for improvement of other cereal
crops. It is a perennial plant, which forms several stalks, and its
pollen can be collected throughout the year when the plant is
maintained in a controlled environment. It can be used as the
pollen parent for wheat crosses for haploid production without
requiring synchronization of flowering dates.
Cultivation
Generally, coix is propagated by seeds. The plant prefers light
(sandy), medium (loamy), and heavy (clay) soils. It prefers
neutral and basic soils but can grow in very acidic soil, and it
188 THE CEREAL GRAINS | Coix: Overview

Name Coixol
Synonyms 6-Methoxy-3H-benzoxazol-2-one

H
N
Molecular Structure O
O O

Molecular Formula C8H7NO3

Molecular Weight 165.15

Name Coixenolide
Synonyms [(2S,3R )-3-[(Z)-hexadec-9-enoyl]oxybutan-2-yl] (E)-octadec-11-enoate

O
H3C
O
Molecular O CH3
Structure H3C
CH3 O
C17566

Molecular
C38H70O4
Formula
Molecular
590.96
Weight
Figure 4 Molecular data and structure of coixol (from www.Chemblink.com) and coixenolide (from Kegg Brite via www.genome.jp).

requires moist soil. It is sown at rate of 610 kg ha1. Seed is
planted 2.5 cm deep, at a spacing of 60  60 cm. One intercul-
tivation, before the plants tiller and cast shade on the ground,
may be necessary. The largest plants result when the plant is
given partial shade although it will tolerate full sun. It adapts to
wet soils and so may be used near water. Sufficient rains in the
early stage of growth and a dry period when grain is setting are
necessary for good yields. Plants respond well to liberal appli-
cations of organic manure. The grain is usually harvested in the
fall when the plant matures and is dried in the sun. Plants are
cut off at base and grain is separated by threshing. Coix yields
depend on varieties cultivated in different countries. Usually,
unhusked grain yields vary from 2.1 to 3.5 tonnes ha1.
Leaf blight, caused by Bipolaris coicis, is one of the most
destructive and major yield-limiting diseases in coix. Plant
response to leaf blight is quantitative rather than qualitative,
varying greatly among cultivars or lines tested. Yield reductions
are primarily associated with decrease in kernel numbers per
plant, thousand kernel weight, and percent kernel ripeness.
Other diseases common in coix are rusts and smuts. Rusts are
caused by Puccinia, while smuts are caused by Tilletia and
Ustilago. Leaf-gall virus and the nematode Meloidogyne incognita
acrita also attack this plant.
Exercises for Revision
Compare and contrast the composition and reported
medicinal claims for coix and other minor grain crops,
such as Amaranthus, buckwheat, and quinoa.
Prepare a map illustrating global production and uses
of coix.
Exercises for Readers to Explore the Topic Further
Write a proposal for the development of a line of nutraceu-
tical ingredients based on coix.
Suggest how traditional food uses of coix could be mod-
ernized and marketed as new products.
Investigate the potential for transfer of medicinally impor-
tant components of coix grain to the high-yielding and
related crop species, maize.
See also: Breeding of Grains: Maize: Breeding; Food Grains and
Well-being: Functional Foods: Dietary Fibers, Prebiotics, Probiotics,
and Synbiotics; Functional Foods: Overview; Genetics of Grains:
Maize: Genetics; The Basics: Taxonomic Classification of Grain
Species; The Cereal Grains: An Overview of the Family of Cereal
Grains Prominent in World Agriculture; Maize: Overview; The
Legumes and Pseudocereals: Amaranth: Overview; Buckwheat:
Overview; Pseudocereals: Overview; Quinoa: Overview.
Further Reading
Ary MB, Richardson M, and Shewry PR (1989) Purification and characterization of an
insect alpha-amylase inhibitor/endochitinase from seeds of Jobs tears (Coix
lachryma-jobi). Biochimica et Biophysica Acta 999: 260266.
Ary MB, Shewry PR, and Richardson M (1988) The amino acid sequence of a cereal
Bowman-Birk type trypsin inhibitor from seeds of Jobs tears (Coix lachryma-jobi
L.). FEBS Letters 229: 111118.
Dante RA, Neto GC, Leite A, Yunes JA, and Arruda P (1999) The DapA gene encoding
the lysine biosynthetic enzyme dihydrodipicolinate synthase from Coix lacryma-
jobi: Cloning, characterization, and expression analysis. Plant Molecular Biology
41: 551561.

Duke JA and Ayensu ES (1985) Medicinal Plants of China. Algonac, MI: Reference
Publications.
Hartwell JL (1971) Plants used against cancer, a survey. Lloydia 34: 386425.
Huang X, Qin J, and Lu S (2014) Kanglaite stimulates anticancer immune responses
and inhibits HepG2 cell transplantation-induced tumor growth. Molecular Medicine
Reports 10: 21532159.
Jain SK and Banerjee DK (1974) Preliminary observations on the ethnobotany of the
genus coix. Economic Botany 28: 3842.
Leite A, Yunes JA, Turcinelli SR, and Arruda P (1992) Cloning and characterization of a
cDNA encoding a sulfur-rich coixin. Plant Molecular Biology 18: 171174.
Li SC (1596) Pen-tsao kang mu (Systematic Pharmacopoeia), China.
Li J and Corke H (1999) Physicochemical properties of normal and waxy Jobs tears
(Coix lachryma-jobi L.) starch. Cereal Chemistry 76: 413416.
Li XH, Huang YQ, Li JS, and Corke H (2001) Characterization of genetic variation and
relationships among Coix germplasm accessions using RAPD markers. Genetic
Resources and Crop Evolution 48: 189194.
List PH and Horhammer L (1979) Hagers Handbuch der Pharmazeutischen Praxis.
Berlin: Springer, vols. 26.
Liu Y, Zhang W, Wang X-J, and Liu S (2014) Antitumor effect of KanglaiteW injection in
human pancreatic cancer xenografts. BMC Complementary and Alternative
Medicine 14: 6, Article Number 228.
Moreau RA, Singh V, and Hicks KB (2001) Comparison of oil and phytosterol levels in
germplasm accessions of corn, teosinte, and Jobs tears. Journal of Agricultural and
Food Chemistry 49: 37933795.
Numata M, Yamamoto A, Moribayashi A, and Yamada H (1994) Antitumor components
isolated from the Chinese herbal medicine Coix lachryma-jobi. Planta Medica
60: 356359.
Schaaffhausen RV (1952) Adlay or Jobs tears: A cereal of potentially greater economic
importance. Economic Botany 6: 216227.
THE CEREAL GRAINS | Coix: Overview 189
Vettore AL, Yunes JA, Neto GC, da Silva MJ, Arruda P, and Leite A (1998) The
molecular and functional characterization of an Opaque2 homologue gene from
Coix and a new classification of plant bZIP proteins. Plant Molecular Biology
36: 249263.
Yamada H, Yanahira S, Kiyohara H, Cyong J-C, and Otsuka Y (1987) Characterization of
anti complementary acidic heteroglycans from the seed of Coix lacryma jobi var ma
yuen. Phytochemistry 26: 32693276.
Yeung HC (1985) Handbook of Chinese Herbs and Formulas. Los Angeles: Institute of
Chinese Medicine.
Zhan YP, Huang XE, Cao J, et al. (2012) Clinical safety and efficacy of KanglaiteW
(Coix Seed Oil) injection combined with chemotherapy in treating
patients with gastric cancer. Asian Pacific Journal of Cancer Prevention
13: 53195321.
Zhang F and Yi Z (1994) Cytological studies of the genus Coix. Natural Science Journal
of Changsha Normal University of Water Resources and Electric Power 9(2):
187191.
Relevant Websites
http://www.fao.org The website of Food and Agriculture Organization of the United
Nations. Search for coix information within the site, such as: http://www.fao.org/ag/
agp/AGPC/doc/Gbase/data/pf000205.htm.
http://www.hort.purdue.edu The website of the center for new crop and plant products
at Purdue University. Search for coix information within the site, such as: https://
www.hort.purdue.edu/newcrop/duke_energy/Coix_lacryma-jobi.html.
http://www.itmonline.org/articles/coix/coix.htm A useful article by Dr. Subhuti
Dharmananda on Coix: Food and Medicine.
 Millet Pearl: Overview
JRN Taylor, University of Pretoria, Hatfield, South Africa
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Pearl millet is quantitatively the most important millet
species.
Pearl millet is uniquely hardy and can be cultivated in hot
and dry conditions.
Pearl millet grain has better nutritional value than most
other cereals.
Many traditional foods and beverages and specialty prod-
ucts are produced from pearl millet.
Pearl millet is becoming widely used in commercial small-
scale food manufacture.
Iron- and zinc-biofortified pearl millet has been developed
for improved nutrition.
Learning Objectives
To provide an introduction to the importance of pearl
millet as a staple food crop.
To provide brief and accurate information about pearl grain
structure, nutritional composition, and traditional food
and beverage uses.
To provide brief and interesting information on the tech-
nologies used to produce traditional foods and beverages
from pearl millet.
To sensitize the reader to the potential of pearl millet as a
global staple food crop in the age of climate change.
Introduction
Pearl millet, commonly known as bulrush millet (Pennisetum
glaucum (L.) R. Br.), also classified as P. typhoides, P. ameri-
canum, or P. spicatum, is a cultivated, small-grain, C4 tropical
cereal grass. Vernacular names include bajra (India), gero
(Nigeria, Hausa language), hegni (Niger, Djerma language),
sanyo (Mali), dukhon (Sudan, Arabic), and mahangu
(Namibia). It is diploid (2n 2x 14) with a large genome
(2450 Mbp). Pearl millet is quantitatively the most important
millet, with a world annual production of some 1525
million tonnes (Mt). It is cultivated mainly in the semiarid
tropics, almost exclusively by subsistence and small-scale
commercial farmers.
This article will examine pearl millet production and agro-
nomic issues, grain structure, chemistry and nutritional
aspects, primary processing technologies, important foods
and beverages produced from pearl millet, pearl millets future
prospects, and research and development challenges.
190 Encyclopedia of Food Grains, Second Edition
Production and Agronomic Issues
Pearl millets great advantage over other cereals is the fact that it
can be and is cultivated in areas of very low rainfall
(300500 mm per year). However, it requires evenly distributed
rainfall during the growing season, and too much rain at flower-
ing can cause a crop failure. Pearl millet can be cultivated in
areas with very high temperatures (daytime temperatures above
30  C), and it germinates well at soil temperatures of 2330  C.
Its importance in food security in dry, marginal agricultural
areas is immense. It is estimated that more than 90 million
people in the semiarid tropics depend on pearl millet food
and income. The major areas of pearl millet cultivation are
India and Western and Central Africa. It is also cultivated in
Eastern and Southern Africa. In Western and Central Africa,
there is a broad band of cultivation from Senegal in the west,
across the Sahel region. It is also cultivated in Sudan, South
Sudan, Eastern Africa, and Southern Africa where there is a
band of cultivation from Angola and northern Namibia, to
Zimbabwe and northern Southern Africa, to Mozambique.
There is also some production in Brazil and the United States.
Figures for pearl millet production are only approximate as
indicated by the wide reported ranges. FAOSTAT data do not
distinguish between the different types of millets. Pearl millet
is generally a subsistence grain crop and hence not commer-
cially traded in quantity. Thus, precise figures are not always
available. Pearl millet accounts for about half of the world
millet crop, which is at least 29 Mt per year. The major pearl
millet grain-producing countries are India,  8.3 Mt per year;
Western and Central Africa,  14.1 Mt, with Niger being the
major producer, followed by Mali, Burkina Faso, Nigeria, and
Senegal; and Eastern and Southern Africa, 1.8 Mt. It is also
increasingly grown as a forage crop and as an ornamental grass.
Relative to other major cereal grains, yields are in general very
low, on average <1 t ha1. This is because pearl millet is grown in
areas of low rainfall and high temperature, often on light, well-
drained soils. Generally, traditional farming practices are used
with very low inputs, that is, no biocides or inorganic fertilizers
and little application of organic fertilizer, and use of traditional
landraces. Pearl millet has, however, much greater potential. In
areas, especially India, where new varieties and hybrids are being
widely cultivated, with some irrigation and using higher-input
agriculture, yields can be well in excess of 2 t ha1.
Postharvest practices also remain backward in many areas, for
example, manual threshing and winnowing of the grain
(Figure 1). Pearl millet can also suffer severe storage losses,
particularly as a result of insects such as the rice moth (Corcyra
cephalonica). The larvae contaminate the grain by producing
webbing (silk) and consume the protein- and fat-rich grain
germs. Control is traditionally affected by putting wood ash in
the storage bin. Today, application of the fumigant phostoxin
(active ingredient phosphine gas) is becoming more widespread.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00011-5

Figure 1 Winnowing of pearl millet in northern Namibia.
Figure 2 Pearl millet grain. Courtesy of Dr L A M Pelembe.
Grain Structure, Chemical Composition,
and Nutritional Value
Grain Structure
Pearl millet grains are tear-shaped to ovoid (Figure 2). They are
up to 2 mm in length, and the 1000-kernel weight is in the
range 315 g (typically  8 g), about one-quarter that of a
wheat grain. The overall grain color varies from pearly white
(hence the name) to yellow, slate gray, brown, or purple.
Individual grains are often not uniformly colored. The antho-
cyanin pigments responsible for grain color are concentrated in
the pericarp.
The general grain structure is essentially the same as other
major tropical cereals, maize and sorghum (Figure 3). The
kernel is naked, that is, it generally threshes free of the hull. A
feature characterizing pearl millet grain structure is the propor-
tionally very large germ and hence relatively smaller endo-
sperm. This impacts the chemical composition of the grain.
THE CEREAL GRAINS | Millet Pearl: Overview 191
Style
Pericarp
Seed coat
Aleurone layer
Corneous
endosperm
Peripheral
endosperm
Floury
endosperm
Scutellar
Epithelium
Scutellum
Embryonic
axis
Hilum
Figure 3 Diagrammatic longitudinal section through a pearl millet
grain. Courtesy of Mr S C Barrion.
The endosperm comprises two major components: the outer
corneous (also referred to as vitreous or hard) endosperm and
the inner floury (also referred to a soft) endosperm. The cells of
the corneous endosperm are filled with an airspace-free con-
tinuous matrix of proteins comprising protein bodies and
matrix protein. It contains few starch granules. In contrast,
the cells of the floury endosperm have airspaces in them and
contain many more starch granules and less protein. The peri-
carp is variable in thickness, dependent on variety, and comp-
osed of three layers: the epicarp, mesocarp, and endocarp. The
mesocarp may contain starch granules such as in sorghum
grain. Beneath the pericarp is a seed coat, which may be
pigmented, and beneath this is the aleurone layer, part of the
endosperm, which is one cell thick. The kernel is enveloped in
a waxy cutin layer, which protects the grain against weathering.
Chemical Composition and Nutritional Value
The general chemical composition and major nutrient contents
of pearl millet grain are given in Table 1. It should be noted
that these values refer to the whole, unprocessed grain. Proces-
sing by milling and malting will affect both the content and
availability of many nutrients.
Energy
Whole pearl millet grain has higher energy content (1720 kJ
100 g1 (db)) than all other cereal grains except maize.
192 THE CEREAL GRAINS | Millet Pearl: Overview

Table 1 Proximate chemical and nutrient composition of pearl millet grain

Proximate composition (db) (g 100 g1)a Minerals (db) (mg 100 g1)a Essential amino acids (g 100 g protein1)b

Food energy (kJ) 16701800 (1720) Ca 41 Isoleucine 3.94.6 (2.3)c

Protein 8.619.4 (14.5) Cl 47 Leucine 9.512.4 (4.4)
Starch 63.178.5 (71.6) Cu 0.5 Lysine 2.83.2 (3.5)
Fat 1.56.8 (5.1) Fe 10.8 Phenylalanine tyrosine 7.19.3 (3.0)
Dietary fiber 8.09.0 (8.5) Mg 125 Threonine 3.34.1 (1.8)
Ash 1.63.6 (2.0) Mn 0.8 Tryptophan 1.41.5 (0.5)
Vitamins (db) (mg 100 g1)a P 373 Valine 4.96.0 (2.9)
Vitamin A (RE)d 24 K 460 Cystine methionine 3.44.4 (1.8)
Thiamin 0.3 Na 17
Riboflavin 0.2 Zn 2.4
Niacin 2.9
Vitamin E 1.9
a
Range and typical values.
b
Typical range.
c
Ideal pattern for 310-year-olds.
d
Retinol equivalents.
Data from Serna-Saldivar S and Rooney LW (1995) Structure and chemistry of sorghum and millets. In: Dendy DAV (ed.) Sorghum and Millets: Chemistry and Technology,
pp. 69124. St Paul, MN: American Association of Cereal Chemists; Klopfenstein CF and Hoseney RC (1995) Nutritional properties of sorghum and the millets. In: Dendy, DAV (ed.)
Sorghum and Millets: Chemistry and Technology, pp. 124168. St Paul, MN: American Association of Cereal Chemists; National Research Council (1996) Lost Crops of Africa, vol. 1.
Washington, DC: National Academies Press; Abdalla AA, El Tinay AH, Mohamed BE, and Abdalla AH (1998) Proximate composition, starch, phytate and mineral contents of 10 pearl
millet genotypes. Food Chemistry 63: 243246.

This is due to its high fat content, a consequence of the rela-
tively large germ.
Carbohydrates
As with all other cereals, the predominant carbohydrate in
pearl millet is starch. The starch content, typically 7172 g
100 g1 (db), is perhaps slightly lower than most other cereals,
due to the fact that the endosperm is proportionally smaller
than other cereals. The percentage amylose in pearl millet
starch is in the range  17.021.5%, which is relatively low
for normal (nonwaxy) starches. Waxy (essentially 100% amy-
lopectin) pearl millet has not been reported. The starch gelati-
nization temperature is in the range 6169  C, typical of
tropical cereal starches. Soluble sugars in pearl millet grain, as
in all sound cereal grains, are low, in the range  1.42.8
100 g1, with the major sugars being sucrose and raffinose.
The dietary fiber content of pearl millet grain is  89 100 g1
(db). The dietary fiber of pearl millet, as with sorghum, is
mainly of the insoluble (water-inextractable) type.
Proteins and amino acids
Pearl millet grain protein content, typically 14.5 g 100 g1
(db), is high for cereal grains. This is because the grain has a
large germ. This, in turn, also affects the balance of Osborne
protein solubility groups and the overall amino acid composi-
tion of the pearl millet protein. Considering the protein frac-
tions, the prolamin (aqueous alcohol-soluble) proteins,
3141% of total protein, are, as in most other cereals, the
major fraction in pearl millet. However, this level is lower
than most other cereals as prolamins are endosperm-specific
proteins. In contrast, the amount of albumin and globulin
(saline-soluble) proteins, 2526% of total proteins, is proba-
bly somewhat higher than in most other cereals.
The prolamin proteins of pearl millet are called pennisetins
and are similar to the prolamins of maize and sorghum, with high
contents of glutamine (2324 g 100 g protein1), alanine (89 g
100 g protein1), and leucine ( 1012 g 100 g protein1), but
relatively low in proline ( 8 g 100 g1 protein1) and cystine
and methionine ( 1 g 100 g protein1 each). Sodium dodecyl
sulfate polyacrylamide gel electrophoresis under reducing condi-
tions shows a major band of Mr  22 kDa and minor bands
of Mr  20 and 10 kDa. Research indicates that, unlike the situa-
tion with sorghum, wet cooking, as occurs in porridge making,
does not substantially reduce the digestibility of pearl millet
protein. This is probably a consequence of the somewhat lower
percentage of prolamin proteins and the low content of cystine.
However, like the other tropical cereals, the prolamins of pearl
millet differ so much from those of wheat that it is considered as
being gluten-free.
The amino acid composition of pearl millet protein
(Table 1) is characterized by a relatively high content (com-
pared with most cereal grains) of the essential amino acid
lysine, in the range 2.83.2 g 100 g protein1. This is a conse-
quence of the relatively high proportion of lysine-rich albumin
and globulins, due to pearl millets large germ. However, the
lysine content of pearl millet does not meet the requirements
of infants, but it is close to that of young school age children.
This emphasizes the importance of supplementation of cereals
in the diet with meat, dairy, or legumes, as sources of lysine.
Pearl millet protein is, as shown, a satisfactory source of the
other indispensible amino acids.
Fats
The fat content of pearl millet grain, typically 5.1 g 100 g1
(db), is high compared with all other cereal grains, except
maize. In the case of both pearl millet and maize, the high fat
content is due to the large germ. As with all other cereals, the
major fatty acids are linoleic acid (C18:2), typically 4345% of
the total; oleic acid (C18:1), 2627% of the total; and palmitic
acid (C16:0), 2021% of the total. The high content of fat and

in particular of polyunsaturated fatty acids means that whole-
grain pearl millet, when milled, is highly subject to deteriora-
tion through oxidative rancidity.
Vitamins
Pearl millet, like other cereal grains, is an important source of B
vitamins (Table 1). These are concentrated in the aleurone
layer and germ. Hence, their content can be adversely affected
by milling. Niacin occurs partially bound to carbohydrate. To
make it fully available, the flour has to be treated with alkali.
This is done by treatment with wood ash in the preparation of
the Western African porridge called to. Because of its high fat
content, pearl millet is a good source of tocopherols (vitamin E),
located mainly in the germ.
Minerals
Pearl millet grain, like other grains, is a good source of most
dietary minerals (Table 1), with the exception of calcium.
Quantitatively, the highest contents are of phosphorus and
potassium. However, the availability of the former and of the
multivalent metal ions is negatively affected by the presence of
phytate (see the succeeding text). The minerals are concen-
trated in the pericarp, aleurone layer, and germ. Hence, their
content can be adversely affected by milling.
Phytochemicals
With regard to phenolic phytochemicals, when compared with
other millets, pearl millet seems to be very low in soluble and
insoluble forms of hydroxybenzoic acids and flavonoids but
contains reasonable levels of both soluble and bound hydro-
xycinnamic acids. However, the available data are very limited,
and it is probable that levels vary greatly with genotype and
cultivation environment.
Antinutrients
There is evidence that pearl millet contains goitrogens. It
appears that the goitrogens are primarily phenolic flavonoid-
type compounds, the C-glycosyl flavones: vitexin, glucosyl
vitexin, and glucosyl orientin. Other phenolic compounds,
such as phloroglucinol, resorcinol, and p-hydroxybenzoic
acid, could also be involved. These compounds apparently
inhibit the deiodination of the hormone thyroxine to its
more active form triiodothyronine. They are concentrated in
the outer layers of the grain and are considerably reduced when
the grain is dehulled during milling. The nutritional signifi-
cance of the goitrogens in pearl millet should not be over-
stated. Although some rural people, for example, in Sudan/
South Sudan and areas of India, who consume pearl millet as a
staple have been found to suffer from goiter, it is probable that
this is because their diet is very restricted and hence deficient in
iodine.
The C-glycosyl flavones contribute to the brown/gray color
of pearl millet and also appear to be responsible for the char-
acteristic musty flavor of damp pearl millet flour, which is
probably of greater significance. Some people describe the
flavor as being mousy or mouse-dropping like. In pearl
millet foods, such as porridges, it can be rather disagreeable
to those not familiar with the food.
THE CEREAL GRAINS | Millet Pearl: Overview 193
Pearl millet, like all grains, contains the phosphorus-
containing compound phytate, myo-inositol hexaphosphate.
Phytate acts as the main phosphorus store in seeds. In grain
foods, it has the undesirable property of binding multivalent
metal cations such as iron, zinc, and calcium and rendering
them biologically unavailable. The level of phytate in pearl
millet grain, in the range 350800 mg 100 g1, is typical of
cereal grains (Table 1). Phytate in pearl millet, as in other
cereal grains, is located in the aleurone layer and germ.
Hence, dehulling the grain during milling substantially
reduces the level of phytate (but also the minerals) in flour.
Malting and fermentation specifically reduce the level of phy-
tate by enzymatically degrading it, freeing the minerals.
It should be noted that contrary to what is written in
some texts, pearl millet, unlike some varieties of sorghum
and finger millet, does not contain condensed tannins
(proanthocyanidins).
Primary Processing
Milling
Pearl millet is milled into flour for porridge and bread making
and into semolina (coarse grits) for couscous making. The
milling process used in northern Namibia is shown in Figure 4
and will be described in detail as it has some unique features.
Dehulling
To improve the palatability and storage quality of the flour, the
bran (pericarp and germ) is generally removed from the grain
first. This process of bran removal is referred to as dehulling.
Cleaned pearl millet
grain
Decorticate
(PRL dehuller)
Lactic acid ferment
(soak overnight)
Partially dry
(solar drying)
Mill
(hammer mill)
Fine flour
Fully dry
(solar dry)
Pack flour
(paper or woven
sacks)
Figure 4 Pearl millet milling process used in northern Namibia.
194 THE CEREAL GRAINS | Millet Pearl: Overview
Figure 5 Small-scale commercial pearl millet mill in northern Namibia.
Foreground hammer mill; background PRL dehuller.
Strictly speaking, decortication is the correct term since the
pearl millet grain does not have a hull.
Figure 5 shows the equipment used in a small-scale com-
mercial pearl millet mill in Namibia. In the background is a
Prairie Research Laboratory (PRL) abrasive-type dehuller.
Today, abrasive-type hullers are manufactured in a number of
countries in Southern and Western Africa. The PRL dehuller
comprises an axle with  12 Carborundum disks mounted on
it, within a cylindrical box. The axle revolves at high speed and
the disks abrade off the bran from the grain. A suction fan
removes the bran. Alternatively, pearl millet dehullers in West-
ern Africa incorporate revolving brushes and screens to remove
and separate the loose bran from the dehulled grain.
The amount of dehulling is determined by the duration of
the dehulling process. Anything between 10% and 30% of the
grain is removed, depending on the desired final color and fat
content of the flour. Dehulling has the effect of reducing the
fiber, fat, and protein contents of the grain due to the removal
of pericarp and germ material. The quality of the protein will
also be adversely affected as the quantity of the lysine-rich germ
proteins is reduced. The starch content is increased slightly.
Steeping
In the Namibian process, the dehulled grain is steeped in
ambient temperature water overnight. This steeping process is
actually a lactic acid fermentation. A lactic acid bacterial cul-
ture, maintained by the process of back slopping, is used to
inoculate the steep water. The culture comprises a portion of
the previous steep, with a high load of suitable lactic acid
bacteria. The purpose of the lactic acid steep seems to be
severalfold. It lightens and brightens the flour. The low pH of
the steep water is responsible for lightening the color and
partially leaching out the brown/gray polyphenolic pigments
of the grain. Goitrogens may also be leached out of the grain.
However, it is probable that water-soluble proteins, vitamins,
and minerals are lost in the steeping process. The steep softens
the grain, facilitating its reduction into a fine flour. The result-
ing flour has an acidic flavor, which consumers prefer.
Figure 6 Pearl millet malt before drying. Courtesy of Dr L A M Pelembe.
Solar drying
The dehulled, steeped grain is partially dried before milling,
and after milling, the flour must be fully dried to a shelf-stable
moisture content of  10%. Solar drying is used for both steps.
The grain or flour is dried on polythene sheets, either directly
on a concrete floor or on long tables made of wooden slats.
Milling
The partially dried, dehulled steeped grain is milled into a fine
flour using a powerful hammer mill. The high moisture con-
tent (3040%) of the grain necessitates relatively powerful
milling. Thus, the process is actually semiwet milling. This
apparently rather inefficient process seems to be necessary in
order to mill the grain into fine flour. In Western Africa, disk
mills are generally used to produce semolina for couscous
making.
Malting
In sub-Saharan Africa, pearl millet is malted to produce malt
for brewing traditional opaque beer and nonalcoholic bever-
ages. Pearl millet malting is almost exclusively done in the
home, although one large-scale commercial malting operation
in Zimbabwe malts pearl millet together with sorghum.
Figure 6 shows pearl millet malt prior to drying. It can be
seen that there is extensive root and shoot growth. The opti-
mum temperature conditions of pearl millet malting are in the
range 2428  C, identical to that for sorghum. The malting
process involves three steps: steeping, germination, and drying.
The steeping time for pearl millet is short, up to 8 h, as the
grain germinates rapidly. Germination is allowed to progress
for 45 days. Like sorghum, pearl millet must be watered
during germination. Drying is at ambient temperature or up
to 50  C in order to conserve malt enzyme activity.
Pearl millet malt quality seems to be somewhat higher than
that of sorghum malt. Diastatic power (total amylase activity)
is similar,  3050 sorghum diastatic units g1. However, the
activity of the b-amylase component is higher. b-Amylase is
the enzyme that produces the fermentable sugar maltose. The
higher b-amylase activity indicates that pearl millet malt could
have potential as partial substitute for barley malt for brewing
lager beer. Free amino nitrogen and extract are also higher in
pearl millet malt than sorghum malt. Of particular significance

is the fact that malting almost completely eliminates the mousy
flavor associated with pearl millet flour. Some nutritional
improvements also occur, in particular improvement in carbo-
hydrate and protein availability. However, against this, there is
a substantial overall loss in mass of grain, 1015%, due to
respiration during germination.
Food and Beverage Production
Worldwide, pearl millet is processed into many different tradi-
tional foods and beverages. Foods include rice and flatbreads
(roti) in India, couscous in Mali and Senegal, doughs such as
fura in Ghana, thick porridges such as to in Western Africa
and bogobe in Southern Africa, thin fermented porridges such
as uji in Eastern Africa, and gruels like ogi in Nigeria. Bever-
ages can be nonalcoholic like oshikundu in Namibia or alco-
holic like the opaque beer of Southern Africa.
Foods
Porridges, doughs, and gruels
In sub-Saharan Africa, pearl millet is most frequently con-
sumed as a porridge. A traditional Pedi (Limpopo Province,
South Africa) recipe to make bogobe bia bupi bja leotsa
(literally porridge of pearl millet) involves cooking 1 kg of
whole-grain pearl millet meal in 1.9 l water. The porridge has
greenish-brown color and a thick, smooth viscous consistency.
Because of the high ratio of starch to water, the porridge gels on
cooling. The flavor is musty, sweet, and bitter. The meal ration
size is 1 kg, which provides some 2205 kJ, 2025% of an
adults recommended dietary allowance. To porridge from
Figure 7 Pearl millet couscous from Senegal.
THE CEREAL GRAINS | Millet Pearl: Overview 195
Western African countries such as Mali and Burkina Faso is
often produced from highly refined flour giving it a gel-like
consistency. It may be cooked with tamarind or lemon or with
wood ash or potash, giving it acidic or alkaline taste, respec-
tively. These porridge products are traditionally eaten cool or
cold with the hand and served with a vegetable relish.
Doughs and gruels made from pearl millet are invariably
lactic acid bacteria-fermented. The fermentation used in fura
dough making has been reported to produce bacteriocins with
inhibitory activity against a wide range of pathogens.
Couscous
In the Sahel region of Western Africa, couscous made from
pearl millet and sorghum is a very popular food (Figure 7).
Couscous making involves processing coarse endosperm par-
ticles (semolina) in repeated steps of agglomerating, steaming,
and sifting to ultimately produce cooked granular particles of
uniform size (Figure 8). The steaming gelatinizes the starch on
the surface of the particles so that they stick together. The
couscous is often served with yogurt, the product being called
thiakri.
Flatbreads
In India, pearl millet is generally consumed as a crisp flatbread,
called roti or chapatti. Figure 9 shows a flowchart for making
roti. A problem in making breads from non-gluten-containing
grains such as pearl millet is that the product tends to break
into pieces. It is notable that in the milling process, the flour is
ground to fine particle size,  43% having a particle size
<75 mm. This increases the surface area of the flour particles
and damages the starch granules, increasing their water absorp-
tion. Both factors increase the cohesiveness of the dough.
196 THE CEREAL GRAINS | Millet Pearl: Overview

Pearl millet grain

Mill
(disk or plate mill)

Whole grain flour

Sieve
(850 mm screen)

Flour (50 g)
+
Warm water (45 ml)

Knead by hand into dough

Flatten and work into

circular sheet
(120 250 mm diameter,
1.3 3.0 mm thick)

Bake (iron or earthenware

pan 300325 C)
top side moistened

Figure 8 Sieving agglomerated pearl millet and sorghum couscous in

Puff
Senegal.
(unmoistened side exposed
to dry heat)

Roti

Variations on the roti-making process, which also alleviate the
problem, can involve cooking part of the flour in water and
mixing it together with the uncooked flour. Cooking gelati-
nizes the starch making it into a binder, as with couscous,
which improves the cohesiveness and elasticity of the dough.
Alternatively, pearl millet flour can be composited with flours
such as gram (mung bean) or other legume flours that give
better cohesiveness or wheat flour that gives elasticity.
Because of the absence of gluten in pearl millet, breads
made from pearl millet flour are unleavened. However, in the
roti-making process, as shown in Figure 9, the flatbread is
exposed on one side to high heat, which puffs it giving the
bread some leavened texture. Puffing takes place as result of
the fact that the moistening water added during baking is very
rapidly turned into steam. The resulting increase in volume
creates an air sac as the steam escapes. Roti is served with a
small amount of hot pickle, dhal, or vegetable sauce.
Beverages
Throughout Africa, sorghum, maize, and millets are used to
produce both nonalcoholic and alcoholic beverages. Invari-
ably, these beverages have undergone a lactic acid fermenta-
tion, giving them a refreshing sour taste. The low pH of the
beverages renders them free from foodborne pathogenic bac-
teria and helps protect them against microbial spoilage.
Figure 9 Roti-making process.
Nonalcoholic
A very popular nonalcoholic fermented beverage in Namibia is
oshikundu made from pearl millet and sorghum malt flour.
Some 200 g of pearl millet flour are thoroughly mixed into
500 ml of boiling water. Then,  100 g of sorghum malt flour
is added. Sorghum malt provides amylase enzymes and lactic
acid bacteria. a-Amylase thins the porridge into gruel by hydro-
lyzing the starch into dextrins, and b-amylase further hydro-
lyzes the dextrins into maltose. The thinning and hydrolysis
actions of the amylases improve food palatability and carbo-
hydrate availability. The lactic acid bacteria use the fermentable
sugars for respiration, producing lactic acid and other flavor
compounds. After the mixture has cooled to room tempera-
ture, 11.5 l of cold water is mixed in. The container is closed
and the mixture is allowed to ferment overnight before con-
sumption. Oshikundu is greenish brown in color and has the
consistency of drinking yogurt. It is slightly effervescent with a
buttery sour taste. A lactic acid content of 0.6% has been
measured.
Opaque beer
In several southern African countries, pearl millet malt is used
to brew traditional, opaque-type beer. Figure 10 shows a
 THE CEREAL GRAINS | Millet Pearl: Overview 197

Pearl millet Ferment in Luke warm Future Prospects and Challenges

malt clay pot water
(3.7 kg) (24 h warm temp) (3.4 l) With the world beginning to experience climatic change, pearl
millet with its unique ability to produce a crop under hot and
dry conditions should become a far more globally important
Decant off
supematant food grain. In the developing world, the need for food and
nutrition security demands that pearl millet increasingly
Supematant Residue becomes a commercially cultivated, traded grain and processed
(8.6 l) into food products, instead of just being a subsistence crop.
The International Crops Research Institute for the Semi-Arid
Bring Tropics (ICRISAT) is leading a project to develop pearl millet
supematent with substantially increased levels of the iron and zinc using
to boil
varietal selection and conventional breeding. Varieties of this
Stir in residue and biofortified pearl millet were released for cultivation in India
boil (1 h) and Africa in 2013. Further, as a result of rapid urbanization
and rising incomes across much of Africa, small-scale commer-
cial enterprises based on making convenience versions of tra-
Hold to gel
ditional pearl millet food and beverage products are becoming
(24 h room temp)
widespread. Related to both nutritional value and commercial
value, the high-energy, fat, and protein content, coupled with
Mash and ferment Pearl millet malt its relatively high level of lysine, is leading to increasing
(24 h warm temp) (1.8 kg) cultivation of pearl millet for poultry feeding, particularly in
Western Africa.
Strain by Strainings Numerous scientific and technological problems still
wringing through (spent grain) require further research and development for pearl millet to
grass bag (4.7 kg) achieve its global potential. These include improving its resis-
tance to a range of plant diseases, creating sustainable seed
Beer production and supply systems, more efficient milling of
(12.3 l) grain to reduce losses, eliminating the mousy aroma in pearl
millet foods, and determining the significance of the reported
Figure 10 Pearl millet opaque beer-making process. goitrogen problem.

flowchart for a Pedi recipe for making bjalwa bja leotsa (lit-
erally beer of pearl millet). The process has some interesting
features. It is noteworthy that only malt is used in the recipe. In
contrast to brewing traditional opaque sorghum beer, no
unmalted grain adjunct is used. In fact, part of the malt is
used as adjunct by cooking it after the first fermentation pro-
cess. The first fermentation process is probably primarily a
lactic acid fermentation, but some fermentable sugar produc-
tion will also take place. As there is no unmalted adjunct in the
recipe, it means that there is a proportionally higher ratio of
amylase enzymes to starch, producing more fermentable
sugars. Coupled with the fact that pearl millet malt intrinsically
contains a higher ratio of b-amylase, this probably accounts for
the fact the pearl millet beer is claimed to be more intoxicating
than sorghum beer. Typical of traditional recipes, more than
one process takes place during a particular step. This is unlike
modern food and beverage manufacturing processes where
there is a system of unit operations. In the mashing and fer-
mentation stage, gelatinized starch is hydrolyzed by the malt
amylase enzymes into fermentable sugars, which are fermented
by both yeast and lactic acid bacteria to produce ethanol,
carbon dioxide, and lactic acid as the main products. The
beer is described as turbid, greenish brown in color, with a
milk like consistency. The flavor is musty and bitter sour, with
a lactic acid content of  1.8%. Pearl millet beer seems to be
much thinner and more acidic than sorghum beer.
Exercises for Revision
Draw the structure of the pearl millet grain, naming the
grain tissues.
Explain the major nutritional advantages of pearl millet
compared with other cereals.
Name the antinutrients present in pearl millet and explain
their nutritional significance.
Draw a flow diagram showing the steps in producing a
traditional pearl millet food of your own choice.
Draw a flow diagram showing the steps in producing a
traditional pearl millet beverage of your own choice.
Explain the science of how textured dry food products can
be produced from pearl millet, despite the fact that it does
not contain gluten.
Exercises for Readers to Explore the Topic Further
There is value in biofortification of pearl millet with other
essential micronutrients than iron and zinc, for example,
provitamin A.
How can the yields of pearl millet in Africa be substantially
improved to help eliminate food insecurity?
The phenolic phytochemicals in a pearl millet need to be far
better characterized across a wide range of genotypes and
198 THE CEREAL GRAINS | Millet Pearl: Overview
individual functional and antinutrient properties
determined.
A more efficient way to mill the pearl millet grain is
required in order to reduce losses and eliminate the
mousy aroma in pearl millet foods.
The real extent of the goitrogen problem in pearl millet
must be established.
See also: Beverages from Grains: Fermentation: Foods and
Nonalcoholic Beverages; The Basics: Grain and Plant Morphology of
Cereals and how characters can be used to identify varieties; Grain:
Morphology of Internal Structure; Taxonomic Classification of Grain
Species; The Cereal Grains: Millet Minor: Overview; Teff: Overview.
Further Reading
Abdalla AA, El Tinay AH, Mohamed BE, and Abdalla AH (1998) Proximate composition,
starch, phytate and mineral contents of 10 pearl millet genotypes. Food Chemistry
63: 243246.
Chandkara A and Shahidi F (2011) Determination of antioxidant activity in free and
hydrolysed fractions of millet grains and characterization of their phenolic profiles
by HPLC-ESI-MSn. Journal of Functional Foods 3: 144158.
Emmambux MN and Taylor JRN (2013) Morphology, physical, chemical and functional
properties of starches from cereals, legumes and tubers cultivated in Africa: A
review. Starch/Starke 65: 715729.
FAOSTAT. Production: Crops. http://faostat.fao.org/ (access 14.02.01).
Germplasm Resources Information Network (GRIN). GRIN Taxonomy for Plants. http://
www.ars-grin.gov/ (assessed 14.02.01).
ICRISAT EXPLOREit. Pearl millet. http://exploreit.icrisat.org/ (accessed 14.01.08).
ICRISAT and FAO (1996) The World Sorghum and Millet Economies. Patancheru, India/
Rome: International Crops Research Institute for the Semi-Arid Tropics/Food and
Agriculture Organization of the United Nations.
Klopfenstein CF and Hoseney RC (1995) Nutritional properties of sorghum and the
millets. In: Dendy DAV (ed.) Sorghum and Millets: Chemistry and Technology,
pp. 124168. St Paul, MN: American Association of Cereal Chemists.
Murty DS and Kumar KA (1995) Traditional uses of sorghum and millets.
In: Dendy DAV (ed.) Sorghum and Millets: Chemistry and Technology,
pp. 185222. St Paul, MN: American Association of Cereal Chemists.
National Research Council (1996) Lost Crops of Africa, vol. 1. Washington, DC:
National Academies Press.
Obilana AB and Manyasa E (2002) Millets. In: Belton PS and Taylor JRN (eds.)
Pseudocereals and Less Common Cereals, pp. 177217. Berlin: Springer.
Quin PJ (1959) Food and Feeding Habits of the Pedi. Johannesburg: Witwatersrand
University Press.
Rai KN, Govindaraj M, and Rao AS (2012) Genetic enhancement of grain iron and zinc
content in pearl millet. Quality Assurance & Safety of Crops and Food 4: 119125.
Serna-Saldivar S and Rooney LW (1995) Structure and chemistry of sorghum and
millets. In: Dendy DAV (ed.) Sorghum and Millets: Chemistry and Technology,
pp. 69124. St Paul, MN: American Association of Cereal Chemists.
Taylor JRN and Emmambux MN (2008a) Millets. In: Arendt E and Dal Bello F (eds.)
Gluten Free Cereal Products and Beverages, pp. 119148. London: Elsevier.
Taylor JRN and Emmambux MN (2008b) Products containing other speciality grains:
Sorghum, the millets and pseudocereals. In: Hamaker BR (ed.) Technology of
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935. Geneva: WHO.
 Millet Minor: Overview
GK Chandi, University of Guelph, Guelph, ON, Canada
GA Annor, University of Ghana, Accra, Ghana
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Minor millets are domesticated and used as food across
different parts of the world.
Millets are generally used for pancakes, flatbreads, porridge,
fermented beverages, boiled grains, gruel, etc.
Minor millets are decorticated before grinding them into
coarse or fine flours or grits of various particle sizes.
Hypoglycemic properties of millets may be attributed to
various factors such as phenolic compounds, soluble fibers,
and starchlipidprotein interactions.
Owing to their potential therapeutic properties and resistance
to severe weather conditions, minor millets have a great
potential to address the health and food security issues.
Learning Objectives
Understand the processing and utilization of minor millets.
Identify the key attributes responsible for nutritional bene-
fits of minor millets.
Introduction
Minor millets are a heterogeneous group of small-grain
cereal species, grown around the world for food and fodder.
Millets comprise a diverse range of species belonging to
different genera, which originated in different parts of the
world. Some of these millets are cultivated by small farmers
in Africa and Asia, while others grow as wild grasses or
weeds (Figure 1, Table 1). Minor millets are highly adapted
to drought and low-fertility soils. The color and structural
features of minor millet grains vary among different genera
(Figure 2, Table 2). Minor millets fall in one of following
genera: Eleusine, Setaria, Paspalum, Panicum, Brachiaria,
Echinochloa, and Digitaria, with some examples including
the following:
1. Finger millet (Eleusine coracana): This millet species is cul-
tivated in more than 25 countries in Africa. It has been
grown in India in the last 4000 years. It is highly adaptable
to higher elevations and hence grown in the Himalayas up
to an altitude of 2300 m. It however has a higher water
requirement than most of other millets.
2. Common or proso millet (Panicum miliaceum): This millet
species was domesticated in China by about 5000 BP and
between 5000 and 4000 BP in Europe. Proso millet is
considered a self-pollinated crop, but natural cross-
pollination may exceed 10%. Proso millet is found high
in mountains up to 3500 m in India. It grows in temperate
climates and is tolerant to wide range of temperatures. It
matures in 6090 days.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00012-7
3. Foxtail or Italian millet (Setaria italica): This millet species is
a native of China and ranks first in its global production. Its
cultivation in Europe dates to about 2000 BP. Although
found in the tropics, it is adapted to temperate regions. It
has a broad range of days of maturity (70120 days). Green
foxtail millet (Setaria viridis) is the wild antecedent of foxtail
millet and has very limited food usage.
4. Kodo millet (Paspalum scrobiculatum): This millet species is
commonly harvested as a wild cereal in tropical Africa and
the Deccan region of India. It was introduced in India almost
3000 years ago. It is self-pollinating, and its florets generally
remain closed during the flowering period. It matures late
compared to other cereals. Sometimes, due to fungal infec-
tions after the rain, kodo millet grains become poisonous.
5. Little millet (Panicum sumatrense): This millet species grows
up to an altitude of 2100 m in India. It is commonly grown
in India and cooked like rice. It is primarily a self-
pollinating crop with about 3.5% cross-pollination. It
matures between 2.5 and 5 months.
Besides the aforementioned millets, there are other minor
millets that are cultivated in restricted geographic regions and
have no importance in world agriculture trade. Some of these
millets include the following:
Barnyard or sawa millet (Echinochloa utilis): This millet species
is grown on a small scale in Japan, China, India, Pakistan,
Nepal, and Korea. Barnyard grasses grow as weeds and
become a nuisance when not managed. It is usually har-
vested within 6 weeks and is grown as a substitute for rice
when paddy crop fails.
Browntop millet (Brachiaria ramosa): This millet species is a
warm-season annual grass and reaches about 1070 cm
height. It can tolerate drought and is known to be cultivated
and/or utilized from wild harvests today, although in very
restricted regions of Kenya, Tanzania, South Africa, and
tropical Asia.
White fonio (Digitaria exilis) and black fonio (Digitaria iburua):
These millet species are commonly eaten by inhabitants of
semiarid regions of West Africa. Their cultivation seems to
have started around 7000 years ago. White fonio is cultivated
throughout much of this region, except Liberia. It is a very
important crop in Southern Mali, Northeastern Nigeria,
extreme Southern Niger, Western Burkina Faso, Eastern
Senegal, and Northern Guinea. Black fonio is found in iso-
lated pockets in the Jos Bauchi Plateau of Nigeria and the
northern parts of Togo and Benin. Fonios are staple crop of
many poor rural communities of these regions and are char-
acterized by a short production time of 10 weeks.
Guinea millet (Brachiaria deflexa): This millet species is also
among minor cereals of Western Africa. However, its cultiva-
tion is confined to the Fouta Djallon Plateau of Guinea and
Sierra Leone. Some cultivars are very early maturing and thus
199
200 THE CEREAL GRAINS | Millet Minor: Overview

Figure 1 Minor millet crops and grain. (a) Foxtail millet, (b) little millet, (c) kodo millet, (d) common millet, (e) barnyard millet, and (f) finger millet.
Reproduced from Balasubramanian S (2013) Processing of millets. In: National Seminar on Recent Advances in Processing, Utilization and
Nutritional Impact of Small Millets. Madurai Symposium, Thamukkam Grounds, Madurai, 13 September 2013.

Table 1 Common name, origin, and occurrence of minor millets

Common name Origin Where found/cultivated

Finger millet (Eleusine coracana) Uganda or neighboring Eastern Africa and in Asia (India and Nepal)
region
Proso (common) millet (Panicum Central and Eastern Asia Ukraine, Kazakhstan, the United States, Argentina, and Australia
miliaceum)
Foxtail millet (Setaria italica) Eastern Asia (China) China, India, Indonesia, the Korean Peninsula, and some parts of southern
Europe
Barnyard millet (Echinochloa utilis) Japan Tropics and subtropics of India
Little millet (Panicum sumatrense) Southeast Asia India, Nepal, Pakistan, Sri Lanka, eastern Indonesia, and western Myanmar
Kodo millet (Paspalum scrobiculatum) India Wild cereal in Western Africa and India

Source: FAO.org.

suited for intercropping cultivation practices with other grain
crops in dry lands under unreliable rainfall.
Nutritional Importance of Millets
consumption of wheat-based products. With total carbohy-
drate and protein contents of about 70% and 12%, respec-
tively, there is no doubt that minor millets contribute
significantly to the energy and protein needs of populations
in regions where they are principal source of food.

Minor millets are good sources of macronutrients and some

Minerals and Vitamins
micronutrients, as shown in Table 3. The gluten-free nature of
these millets makes them a reasonable alternative for people Minor millets especially finger millets are important sources of
suffering from celiac disease and allergies that result from the minerals especially calcium relative to other cereals. With
 THE CEREAL GRAINS | Millet Minor: Overview 201

Figure 2 Minor millet grains. Reproduced from Balasubramanian S (2013) Processing of millets. In: National Seminar on Recent Advances in
Processing, Utilization and Nutritional Impact of Small Millets. Madurai Symposium, Thamukkam Grounds, Madurai, 13 September 2013.

Table 2 Structural comparison of some minor millets

Plant height
Millet (cm) Seed color Seed shape Seed size (mm) Thousand kernel weight (g, 12% mb)

Finger millet 40100 Yellow, white, red, brown, violet Globose 12 2.6
Proso millet 30100 White cream, yellow, orange, red, black, Oval 3
2 4.77.2
to brown
Foxtail millet 0.5200 Yellow, pale yellow Elliptical 23 1.86
Barnyard 50100 Yellow, red, brown Globose 1.3 4.17
millet
Little millet 3090 White, yellow, red, brown Elliptical 2.3
1.6
Kodo millet  55 Light red to dark gray

Source: Upadhyaya HD, Dwivedi SL, Singh SK, Singh S, Vetriventhan M, and Sharma S (2014) Forming core collections in Barnyard, Kodo, and Little Millets using morph agronomic
descriptors. Crop Science 54: 26732682 and FAO.org.
202 THE CEREAL GRAINS | Millet Minor: Overview

Table 3 Nutrient profile of minor millets (per 100 g edible portion)

Parameter Finger millet Proso millet Foxtail millet Little millet Kodo millet Barnyard millet

Moisture (g) 13.1 11.9 11.2 11.5 12.8 11.9

Protein (N 6.25) (g) 7.3 12.5 12.3 7.7 8.3 6.2
Fat (g) 1.3 1.1 4.3 4.7 1.4 2.2
Minerals (g) 2.7 1.9 3.3 1.5 2.6 4.4
Dietary fiber (g) 11.5 2.4 2.5 25 2.08
Carbohydrates (g) 72.0 70.4 60.9 67 65.9 65.5
Energy (kJ) 1372 1426 1384 1426 1284 1284
Vitamins
Beta-carotene (mg) 42 0 32 0 0 0
Thiamine (mg) 0.42 0.20 0.59 0.30 0.303 0.33
Riboflavin (mg) 0.19 0.18 0.11 0.09 0.09 0.10
Niacin (mg) 1.1 2.3 3.2 3.2 2.0 4.2
Total B6 (mg)
Folic acid (mg)
Free 5.2 4.2 2.2 7.4
Total 18.3 15.0 9.0 23.1
Vitamin C (mg) 0 0 0 0 0 0
Choline (mg) 748
Minerals and trace elements
Calcium (mg) 344 14 31 17 27 20
Phosphorus (mg) 283 206 290 220 188 280
Iron (mg) 3.9 0.8 2.8 9.3 0.5 5.0
Magnesium (mg) 137 153 81 133 147 82
Sodium (mg) 11 8.2 4.6 8.1 4.6
Potassium (mg) 408 113 250 129 144
Copper (mg) 0.47 1.60 1.40 1.00 1.60 0.60
Manganese (mg) 5.49 0.60 0.60 0.68 1.10 0.96
Molybdenum (mg) 0.102 0.70 0.016
Zinc (mg) 2.3 1.4 2.4 3.7 0.7 3.0
Chromium (mg) 0.028 0.020 0.030 0.180 0.020 0.090
Sulfur (mg) 160 157 171 149 136
Chlorine (mg) 44 19 37 13 11
Essential amino acids (mg/16 g N protein)
Arginine 300 290 220 250 270
Histidine 130 110 130 120 120
Lysine 220 190 140 110 150
Tryptophan 100 050 060 060 050
Phenylalanine 310 310 420 330 430
Tyrosine 220
Methionine 210 160 180 180 180
Cystine 140 100 090 110
Threonine 240 150 190 190 200
Leucine 690 760 1040 760 650
Isoleucine 400 410 480 370 360
Valine 480 410 430 350 410

Source: Shobana S, Krishnaswamy K, Sudha V, et al. (2012) Finger millet (ragi, Eleusine coracana L.): A review of its nutritional properties, processing, and plausible health benefits.
Advances in Food and Nutrition Research 69: 139; Gopalan C, Rama Sastri BV, and Balasubramanian SC (2009) Nutritive Value of Indian Foods. Hyderabad, India: National Institute of
Nutrition, Indian Council of Medical Research. Geervani P and Eggum BO (1989) Nutrient composition and protein quality of minor millets. Plant Foods for Human Nutrition 39(2): 201208.

calcium levels of about 344 mg/100 g, finger millet arguably
has the highest calcium content when compared with other
cereals. Using proton-induced x-ray emission spectrometry,
Kruger et al. confirmed the high calcium content of finger
millet and also showed that calcium was distributed through-
out the endosperm where phytate levels were low. Phosphorus,
iron, magnesium, potassium, sulfur, and zinc, for example, are
all found in considerable amounts in minor millets. In terms of
vitamins, some minor millets have been found to have low
amounts of carotenoids thiamine, riboflavin, niacin, and
choline. Finger and foxtail millets have been reported to con-
tain 42 mg/100 g and 32 mg/100 g carotenoid, respectively
(Table 3).
Fat
Fat in minor millets, as in other cereals, is mainly composed of
unsaturated fatty acids. Oleic and linoleic acids are the main
unsaturated fatty acids in minor millets, with palmitic acid as
the main saturated fatty acid. Depending on the species of

millet palmitic, oleic and linoleic acids can range from 6% to
23%, 20% to 66%, and 22% to 65%, respectively, of the total
lipid content of the millet flours. Though these unsaturated
fatty acids may have health benefits, they are also responsible
for the rancidity that is normally observed in stored millet
flours with relatively high fat contents.
Fiber
The dietary fiber contents of minor millets also confer some
health benefits when they are consumed. Total dietary fiber
values of about 27%, 20%, and 30% of grain weight have been
reported for barnyard, kodo, and little millets, respectively, out
of which 0.59%, 1.94%, and 1.33% are soluble fiber,
respectively.
Amino Acids
Just like most of the cereals, the minor millets are deficient in
lysine, which is usually complemented when mixed with
legumes.
Phytochemicals
The pericarp of minor millets is rich in several phenolic
phytochemicals, which are associated with various health ben-
efits. The inhibitory effects of finger millet phenolic com-
pounds on a-glucosidase and pancreatic a-amylase have been
mentioned. It is thought that these millet phenolic compounds
noncompetitively inhibit carbohydrate-hydrolyzing enzymes.
Millet phenolic compounds have also been suggested to delay
or interfere with the absorption of dietary carbohydrates in the
small intestine resulting in the suppression of postprandial
glucose spikes. A comprehensive review of phytochemicals in
minor millets and their health benefits has been reported by
Taylor et al. and Taylor and Duodu.
Hypoglycemic Properties
The hypoglycemic property of millets and its importance in the
management of type II diabetes were reported as far back as
1957. Since then, several other scientists have also reported on
the lowering effects the regular consumption of minor millets
has on postprandial blood glucose levels. It will also be of
interest to understand the factors that contribute to the hypo-
glycemic property of millets to help maintain this property
during processing and also to develop hypoglycemic food
products. Some attempts have been made in this regard.
Role of Fiber
Dietary fiber in millets has been mentioned as one of the
reasons for its hypoglycemic property. With millets containing
about 2% soluble and 30% insoluble fiber contents, their role
in hypoglycemic property of millets cannot be overlooked.
Fiber reduces the glycemic response of foods by increasing
their viscosity, especially the soluble fiber. A considerable
increase in soluble fiber content of finger millet is observed
after decortication, though total dietary fiber decreases. This
THE CEREAL GRAINS | Millet Minor: Overview 203
increase in the soluble fiber content has special nutritional
significance due to its physiological advantages in terms of
hypoglycemic and hypocholesterolemic characteristics.
StarchProtein Interactions
Some researchers have attributed the hypoglycemic property of
finger millet to their rigid starch granule architecture. This
rigidity slows down the rate at which the millet starch granules
are hydrolyzed by starch digestive enzymes. The effects of
protein-encapsulated granule matrix have been suggested as
part of the reason for the hypoglycemic property of kodo
millet. Some protein fractions glue protein bodies into a matrix
surrounding starch granules, which then acts as a barrier to
amylases. The removal of protein from kodo millet flour
clearly results in a significant increase in its starch hydrolysis.
StarchLipid Interactions
The presence of starchlipid complexes, particularly lipid com-
plexes with millet amylose, also influences the susceptibility of
millet starch to enzymatic degradation. Annor et al. reported
significant effects of starchlipid complexes on the starch
hydrolysis rates of millets. Foxtail, little, barnyard, kodo, and
proso millets still contain a considerable amount of lipids even
after decortication. The residual lipids in the endosperm are
still sufficient to result in significant reductions in starch hydro-
lysis rates of the decorticated millet flour.
Fatty Acid Profile
In addition to starchlipid complexes influencing the glycemic
properties of millets, the type of lipid and their amounts pre-
sent in millets also significantly contribute to their
hypoglycemic property. The main fatty acids present in millets
in no particular order are palmitic, oleic, and linoleic acids.
These three fatty acids form more than 80% of the total fatty
acids present in millets. These fatty acids form complexes with
millet starches and thereby reduce the rate at which the result-
ing starchlipid complexes are hydrolyzed by digestive
enzymes. The unsaturated fatty acids form more complexes
than the saturated fatty acids. With regard to their effects on
starch hydrolysis rates, all the fatty acids significantly reduce
the millet starch hydrolysis rates and their expected glycemic
index. Starcholeic acid complex are the most stable against
amylolysis. The amounts of these fatty acids present in the
respective millet flour samples also significantly influence the
expected glycemic index of millet flours. The greater the quan-
tity of a particular fatty acid that is present in the millet flour,
the more it reduces the starch hydrolysis rates and their result-
ing expected glycemic index. It is therefore important to retain
the fatty acids present in millet flours when they are being
processed to utilize their beneficial hypoglycemic property.
Apart from hypoglycemic benefits, the regular consump-
tion of minor millets has been associated to a myriad of health
benefits. Germinated finger millet flour has been reported to
improve hemoglobin status of school children in Tanzania and
also have antimicrobial, antifungal, antioxidant free radical-
scavenging, antiulcerative, cholesterol-lowering, and wound-
healing properties.
204 THE CEREAL GRAINS | Millet Minor: Overview

Storage of Grains
Minor millets are important food crops of a large group of
people living in rural, tribal, and hilly areas across the globe.
About 90% of millets produced are used as food, and the rest of
it is used as fodder or for medicinal purposes. The millet grains
are stored either as unthreshed along with their ear heads or as
threshed and clean grains. The unthreshed grains are normally
heaped on the ground and covered with straw. The threshed
and clean grains are packed in jute sacks and stacked in piles on
the floor or wooden platforms. Some farmers also utilize silos
or bins to store grains when needed. The larger storage struc-
tures are built with metal, wood, stone, or bamboo. In the
absence of silos, the grains are stored in pits lined with straw.
Once the pit is full, the grains are covered with straw and soil or
dung. Clay pots are also sometimes used to store the millet
grains. Occasionally, grains are exposed to sunshine to dry and
also to prevent them from insect infestation.
Postharvest Processing
faster cooking of the grain. For food purposes, postharvest
processing of millets depends on the cultural preferences and
traditions.
Traditional Processing
For the small-headed grains, husk (hull) is usually removed by
hand pounding and less often by mechanical dehusking or
polishing units. Grain is sometimes moistened before pound-
ing. This is done by adding about 10% water to loosen the
fibrous bran and also to separate the germ and the endosperm,
if desired. The traditional processing of minor millets involves,
in no particular order, dehulling, winnowing, washing, drying,
milling, and sieving. Local small-scale mills are usually based
on crude stone mortar and pestle or one horizontal hand
millstone rotating on another. Both types of mills deliver
whole meal. A wooden pestle and mortar (Figure. 4) is also
commonly used in which by dampening, pounding,
winnowing, and pounding again, millet is separated into fine
and coarse semolina, flour, and bran.

Figure 3 depicts the major steps involved in processing of the

millets. The millet kernels are smaller as compared with the
size of majority of the cereal grains. The color of the millets is
confined to the pericarp, which is strongly adhered to the
endosperm. Millet milling is done to remove the hulls and
bran (pericarp) of the grain. It lightens the color, improves
the overall acceptability and nutritional quality, and leads to
Mechanical Processing
Treatments such as parboiling, steaming, and drying are occa-
sionally applied prior to mechanical milling of millets to facil-
itate complete removal of the husk. Parboiling is reported to
help in dehusking kodo millet and to eliminate the stickiness
in cooked finger millet porridge. Parboiling is basically the

HARVESTING
Stems/Panicles/long straws/weeds/leaves

THRESHING AND WINNOWING

Husk/stems/chaffs/weeds
Spikelets/long straws/chaffs

STORAGE
Spikelets/same sized weeds

SOURING
POPPING MALTING
MILLING AND POUNDING (Lactic acid
fermentation)
Grain/chaff/small weeds Husk/chaff/weeds
Tempering Steep
Conditioning Cooking

Heating Germinate Mashing

Dehusking/Debranning
Roller Milling Grinding and Sieving Kiln Straining
Popped grains
Grinding
Polished grains Alcoholic fermentation
Adjuncts for brewing
Fully refined flour Flour
Opaque Beer

Figure 3 Postharvest processing of minor millets.


Figure 4 Women husking fonio millet. Reproduced from Gari JA (2001)
Review of African millet diversity. IPGRI/IFAD, Bamako, Mali, 1922
November 2001, www.FAO.org.
process of partial cooking the grain along with its husk or bran.
The resulting product is dried and decorticated.
Decortication
Decortication improves the biological availability of nutrients
and consumer acceptability of minor millets. It reduces total
protein and lysine, but it also improves the utilization of the
remaining protein. Decortication is sometimes accomplished
by using rice dehullers or other abrasive dehullers. Technically,
there are three types of decortication techniques that can be
employed to minor millets:
1. Abrasive decorticators: Abrasive decorticators abrade away
the fibrous pericarp (Figure 5). If all parts of the grains
could be abraded away at the same rate, abrasive decortica-
tion would be an efficient way of removing the pericarp.
However, different parts of individual grains are abraded
away at different rates, and there is some loss of endosperm
(particularly from damaged grains) even when the grain is
only lightly abraded.
2. Rubbing technique: For this technique, decortication is
achieved by a steel rotor, rotating the grain mass within a
generally cylindrical chamber. When the grain is properly
tempered, the pericarp is rubbed off by the movement of
one seed against another. However, when the grain is too
dry, abrasion of the internal components of the mill
becomes severe. The hulls and the endosperm fragments
are separated in a cyclone, and the endosperm particles are
milled in a proprietary mill.
3. Metal friction machines: This is when attrition mills are used.
Attrition usually means particles getting smaller due to their
THE CEREAL GRAINS | Millet Minor: Overview 205
Figure 5 Tangential abrasive dehulling device. Reproduced from
Balasubramanian S (2013) Processing of millets. In: National Seminar on
Recent Advances in Processing, Utilization and Nutritional impact of
small millets. Madurai Symposium, Thamukkam Grounds, Madurai, 13
September 2013.
corners or surface irregularities being knocked off. These
techniques are used only to a limited extent at commercial
scale.
Milling
After dehulling, the material separated from the endosperm by
aspiration comprises the hull, bran, germ, and also a portion of
the endosperm in the form of broken grains and powder,
which is used as poultry feed. The types of mills used for
pulverizing minor millets are hammer mills, burr mills, and
plate or disk mills. Wet milling is not performed for the minor
millets. Most of the minor millet processing involves semiwet
or dry milling techniques. Hammer mills are probably the
cheapest and simplest mills used for reducing the particle size
of the decorticated grains. These mills are available in different
sizes depending on the size of operations. They consist of
hammers installed on the rotor rotating rapidly in an enclosed
casing. The outlet is covered by a screen that determines the
particle size of the flour. Overheating and moisture built up
during the hammer milling result in caking of the flour and
cause reduction in the throughput of the process. Roller flour
milling is rarely used for minor millets as it is designed for
wheat milling. If roller mills are used for separating the endo-
sperm from the offal, the particle size is usually reduced in
roller mills with smooth rollers. In cereals, the functional
properties of flours are defined by the particle size distribution
and damaged starch contents. However, there is not much
research to evaluate the differences in the functionality of the
millet flours owing to different milling procedures. Also, there
206 THE CEREAL GRAINS | Millet Minor: Overview
appears to be a difference between the flour produced by the
wooden pestle and mortar and that produced by mechanical
methods. The former seems to have better nutritional qualities.
The biggest challenge is to develop a dehulling and milling
technology according to specific physical characteristics of mil-
let grains that can yield flour with uniform quality. It is impor-
tant to note that some efforts have been made in this regard.
The growth of this industry will depend upon constant
demand of millet-based products and regular feeding of the
mills with millet grains, which have been properly harvested
and stored in good conditions.
Recent Advancements
A new eco-friendly and energy-efficient continuous-type CIAE-
Millet mill (models I and II) (Figure 6) has been designed and
developed for all kinds of minor millets. It has provisions to
adjust the clearance between the dehusking surfaces to suit the
different sizes of minor millets. The dehusking efficiency of the
machine is about 95%. The separation of the husk is simulta-
neous with the suction arrangement and cyclone separator.
Another compression-based and shear-based machine has
been patented in India to dehull the preconditioned grains.
The hulls are removed using aspirator. The dehulled grains are
separated using conventional grader and hot air-dried to 10%
moisture content (dry basis). Dried dehulled grains are then
polished in abrasive-type polishers where bran and germ are
removed from the grain. The germ and bran are separated by
screen. Then, the polished endosperm is ground in a burr mill
to obtain refined flour of the required particle size. In another
patent filed by the Indian Agriculture Research Institute
Figure 6 CIAE-Millet mill (models I and II). Reproduced from Balasubramanian S (2013) Processing of millets. In: National Seminar on Recent
Advances in Processing, Utilization and Nutritional Impact of Small Millets. Madurai Symposium, Thamukkam Grounds, Madurai, 13 September 2013.
(2011), a thresher-cum-pearler for finger millet and barnyard
millet has been designed and developed. It works on the prin-
ciple of impact and shear on the grains. The threshing drum is
provided with six leather flats, which provide gentle impact
and shear and a mild rubbing action for threshing, dehusking,
pearling, and polishing of the grains. A sieve assembly is also
attached with different operations. The design is simple and
easily adaptable in rural areas for efficient postharvest proces-
sing of these crops.
Utilization of Minor Millets
In some regions, minor millets remain cultivated only on a
small scale but are culturally important for particular food
stuffs, such as ritual breads made from browntop millet in
India. Some varieties of proso and foxtail millets were histor-
ically used to prepare noodles in Northwestern China 4000
years ago. In Europe and Asia, green foxtail grains are eaten as
boiled, roasted, or ground into flour. The Chinese call this
simply gao wei cao (dog tail grass) to distinguish it from culti-
vated foxtail millet. Yellow bristle grass (S. pallidifusca) grains
are used as fodder in Nepal. Plains bristle grass (S. macrosta-
chya) is a pre-Columbian and historic Mexican cereal. The
grains are toasted, ground, and eaten as gruel. Yellow foxtail
millet (S. glauca) has been observed to be parboiled in South
India.
In general, foods prepared with minor millets can be clas-
sified into two broad categories whole-grain millet foods and
flour foods prepared after grinding of millets. Some of the
popular millet foods of each category (Figure 7) are described
in the succeeding text.

Figure 7 Foods prepared with minor millets in India. Source: Millet
Network of India Deccan Development Society FIAN, India.
The Whole-Grain Millet Foods
These are prepared with millet grains with or without milling
and need little preparation before cooking. Most of the millet
grain foods are prepared by boiling with water:
1. Pudding: Millet pudding is prepared by boiling proso millet
grains with water or milk until they become soft and then
mixed with sugar or jaggery and garnished with nuts and
raisins. It is usually prepared on festivals or special
occasions.
2. Boiled grains: The preparation is similar to rice. The polished
barnyard millet grains are washed and boiled in hot water
over a strong fire for about 6 min. and stirred occasionally.
After draining the excess water, the grains are steamed over
a weak fire for 3 min. The cooked grain complements veg-
etable stew or yogurt.
3. Porridge: Millet porridge is prepared with whole or broken
grains of kodo, foxtail, or finger millets. Sometimes, the
grains are sprouted before boiling with water. Sprouting
greatly improves the carbohydrate digestibility and palat-
ability. This is of particular importance in weaning foods.
4. Millet gruel: This is prepared by boiling broken kodo,
foxtail, or finger millet grains with excess water, salt, and
pepper until the grains are tender and fully cooked.
5. Popped grains: Proso or finger millet grains are sprayed with
water and tempered to equilibrate the moisture. The tem-
pered grains are popped at high temperature for short time.
They are used as snacks or adjuncts in brewing. The popped
grains may be ground into flour and blended with legume
flours. These flours are used in weaning and supplementary
foods.
6. Dumpling: It is prepared by using a composite mix of the
broken proso millet grains and flour. Grains and flour are
mixed in a 1:4 proportion and cooked in boiling water for
3 min. The cooked batter is well kneaded with a wooden
stick and hand-shaped into balls.
7. Stir fried millets: This seems to be a new variation of millet
foods that has appeared recently. The broken grains of
barnyard millet grains are washed in water and added to
the stir-fried vegetables with spices and boiled together.
THE CEREAL GRAINS | Millet Minor: Overview 207
The Millet Flour Foods
These are mostly prepared by baking or frying with vegetable
oil:
1. Unleavened bread (roti): In some regions of south India,
browntop millet is particularly preferred for making tradi-
tional breads. The flour is kneaded with water and mixed
with chopped onion. Chopped green chilies, broken
groundnut, and salt are added to taste. Dough is shaped
into a flat round pancake by hand or rolling pin and then
baked with a spoonful of oil in a frying pan on both sides.
Finger millet is also used for making rotis.
2. Leavened thin pancake: It is common in south India by the
name of dosai. Finger millet flour is mixed with water to
form a thin batter, which is spread on a frying pan and
baked with a spoonful of vegetable oil for about 4 min.
Dosai is normally served with a vegetable stew. Another
kind of pancake is injera, which is a staple food of Ethiopia.
It is commonly prepared with teff and to some extent with
millets. It is 50 cm diameter leavened pancake with a
honeycomb-like texture. Dough is partly cooked to gelati-
nize the starch; the carbon dioxide produced by the lactic
acid fermentation leavens the injera on baking as it escapes.
Additionally, the low pH protects these foods against the
growth of pathogenic bacteria.
3. Snack foods: The common millet snack foods in India are
nippattu, chakkulli, haralu, kodubale, and kadubu. These are
deep-fried snack foods prepared by mixing flour of brown-
top millet or finger millet with black gram flour or wheat
flour in various proportions. Fried Bengal gram and
groundnut are mixed while preparing the dough for
nippattu.
4. Millet sweet dish: Flour of browntop millet is mixed with
water to make dough. A small quantity of this dough is
taken and flattened using a rolling pin and then stuffed
with sweet ingredients and deep-fried in groundnut oil
until golden brown. Traditionally, it is known as kadubu
in India.
Apart from the food uses, minor millets also have been used for
therapeutic reasons. For example, bitter seeds of Liebmanns
bristle grass (S. liebmannii) are used to treat toothache in Mex-
ico. Palm grass (S. palmifolia) is a part of a religious festival of
new rice harvest in the Philippines. Shoots are cooked as green
vegetable in New Guinea. Sections of the stem of the young
plant are steamed in a bamboo tube and then eaten. The
Topnaar people of Namibia use seeds of bristly foxtail (S.
verticillata) in porridge and also to brew beer. It has been
used as fodder in Okinawa and the Ryukyu Islands (Japan),
while in India, the grains are eaten. Besides the traditional
foods, there are lots of novel foods being developed and
researched worldwide, for example, convenience mixes, com-
posite flours, weaning foods, fermented and germinated foods
and gruels, and alcoholic and nonalcoholic beverages.
Millets are largely consumed in rural areas and are com-
monly known as poor mans grains. However, in recent years,
local crop patterns have completely changed due to the popu-
larity of other crops like wheat and rice, which brings better
monetary returns to the farmers. The global availability of
208 THE CEREAL GRAINS | Millet Minor: Overview
inexpensive foods prepared with other cereals like wheat
breads and biscuits has made minor crops like millets lose
their significance even in the local regions where they are
produced. Marketing millets as nutraceutical grains, while
highlighting their key health benefits to create awareness
among consumers, could trigger demand of the millet-based
products. Support from the local governments at financial and
policy framework levels and involvement of cooperatives may
help establish millet processing industry as a model in selected
sites, which can be replicated in other parts of the world. Minor
millets are early maturing and can be cultivated under severe
drought conditions. Thus, there exists a great potential of
exploiting these underutilized genetic germplasms to address
the current and future issues of health and global food security.
Exercises for Revision
Give a few examples of minor millets and their genera.
What is the nutritional significance of polyphenols in
minor millets?
What are the factors responsible for the hypoglycemic prop-
erty of minor millets?
List a few food products prepared from minor millet flour.
Exercises for Readers to Explore the Topic Further
Find out the production and consumption trends of millets
in the last three decades.
Investigate the recent advances in processing of minor mil-
let grains.
Explore how minor millets could be a solution to global
food security issues.
Determine how the molecular structure of minor millet
starches influences their hypoglycemic properties.
Acknowledgments
We gratefully acknowledge the late Dr. Koushik Seetharaman
for the inspiration to write this article. We also appreciate the
support from Canadian International Food Security Research
Fund International Development Research Center and the
Department of Foreign Affairs, Trade and Development, Gov-
ernment of Canada.
See also: Agronomy of Grain Growing: Sorghum: Production and
Improvement Practices; Food Grains and the Consumer: Cultural
Differences in Processing and Consumption; Grain Harvest,
Storage and Transport: The Nature, Causes, and Control of Grain
Diseases in the Major Cereal Species; Grains Around the World:
Grain Production and Consumption: Africa; Non-wheat Foods:
Sorghum: Utilization; Proteins: Protein Synthesis and Deposition;
Scientific Organizations Related to Grains: Research
Organizations of the World: CGIAR; The Basics: Grain: Morphology of
Internal Structure; Taxonomic Classification of Grain Species; The
Cereal Grains: An Overview of the Family of Cereal Grains Prominent
in World Agriculture; Cereals: Domestication of the Cereal Grains;
Millet Pearl: Overview; Teff: Overview; The Legumes and
Pseudocereals: Amaranth: Overview; Wheat-Based Foods:
Cereals: Overview of Uses: Accent on Wheat Grain.
Further Reading
Annor GA (2013) Millet Starches: Structural Characteristics and Glycemic Attributes.
Doctoral dissertation, Canada: University of Guelph.
Annor GA, Marcone M, Bertoft E, and Seetharaman K (2013) In vitro starch digestibility
and expected glycemic index of kodo millet (Paspalum scrobiculatum) as affected by
starchproteinlipid interactions. Cereal Chemistry 90(3): 211217.
Balasubramanian S (2013) Processing of millets. National Seminar on Recent Advances
in Processing, Utilization and Nutritional Impact of Small Millets. Madurai
Symposium, Thamukkam Grounds, Madurai, 13 September 2013.
Bora P (2014) Nutritional Properties of Different Millet Types and Their Selected
Products. Masters dissertation, Canada: University of Guelph.
Devi PB, Vijayabharathi R, Sathyabama S, Malleshi NG, and Priyadarisini VB (2011)
Health benefits of finger millet (Eleusine coracana L.) polyphenols and dietary fiber:
A review. Journal of Food Science and Technology 51(6): 10211040. http://dx.doi.
org/10.1007/s13197-011-0584-9.
Geervani P and Eggum BO (1989) Nutrient composition and protein quality of minor
millets. Plant Foods for Human Nutrition 39(2): 201208.
Kruger J, Pina-Vardas C, Vinis-Ndimba R, and Taylor JRN (2014) Visualisation of the
distribution of minerals in red non-tannin finger millet using PIXE microanalysis.
Journal of Cereal Science 60: 13.
Rajasekaran NS, Nithya M, Rose C, and Chandra TS (2004) The effect of finger millet
feeding on the early responses during the process of wound healing in diabetic rats.
Biochimica et Biophysica Acta 1689: 190201.
Ramanathan MK and Gopalan C (1957) Effect of different cereals on blood sugar levels.
Indian Journal of Medical Research 45: 255262.
Shobana S, Krishnaswamy K, Sudha V, et al. (2012) Finger millet (ragi, Eleusine
coracana L.): A review of its nutritional properties, processing, and plausible health
benefits. Advances in Food and Nutrition Research 69: 139.
Sripriya G, Chandrasekharan K, Murthy VS, and Chandra TS (1996) ESR spectroscopic
studies on free radical quenching action of finger millet (Eleusine coracana). Food
Chemistry 57(4): 537540.
Taylor JRN, Belton PS, Beta T, and Duodu KG (2014) Increasing the utilisation of
sorghum, millets and pseudocereals: Developments in the science of their phenolic
phytochemicals, biofortification and protein functionality. Journal of Cereal Science
59(3): 257275.
Taylor JRN and Duodu KG (2014) Effects of processing sorghum and millets on their
phenolic phytochemicals and the implications of this to the health-enhancing
properties of sorghum and millet food and beverage products. Journal of the
Science of Food and Agriculture 95(2): 225237. http://dx.doi.org/10.1002/
jsfa.6713.
Tovey FI, Jayaraj AP, and Clark CG (1975) The possibility of dietary protective factors in
duodenal ulcer. Postgraduate Medical Journal 51: 366372.
Upadhyaya HD, Dwivedi SL, Singh SK, Singh S, Vetriventhan M, and Sharma S (2014)
Forming core collections in Barnyard, Kodo, and Little Millets using morph
agronomic descriptors. Crop Science 54: 26732682.
 Teff: Overview
G Bultosa, Haramaya University, Dire Dawa, Ethiopia; Botswana College of Agriculture, Gaborone, Botswana
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Teff crop description, origin, cultivation, and production.
Teff grain production constraints and progress of
improvement.
Teff grain morphology, anatomical structure, composition,
and physicochemical and functional properties of the
major constituents of teff grain.
Teff grain nutrient merits, always consumed as whole-grain,
gluten-free alternative food for celiac patients, high in min-
eral nutrients particularly in iron with well-balanced amino
acids.
Processing of teff grain flour to injera (lactic acid bacteria
and yeast-fermented, soft, spongy, sour, circular flatbread,
staple food for majority of Ethiopians).
Prospect for future teff productivity improvement and
utilization and diversification of teff products.
Learning Objectives
Achieve understanding on teff origin, description, grain
production, and constraints associated to grain production
systems.
Understand and apply the knowledge of teff grain mor-
phology, anatomical structures, nutrient compositions,
and grain component functionalities for processing into
various food products such as for gluten-free and whole-
grain foods.
Achieve understanding on teff injera processing stages,
nature of fermentation involved, characteristic of injera for
global consumers, and highlights on other traditional teff
grain food products.
See teff as a traditional Ethiopian grain, strongly linked to
national dietary culture, but with nutritional and sensory
benefits that could lead to product development in new
markets.
Introduction
Teff (Eragrostis tef (Zucc.) Trotter), today generally written as
tef, is a tropical cereal. Teff cultivation and usage as human
food are mostly confined to Ethiopia. This article highlights the
origin, cultivation of the teff plant, and production and storage
of the teff grain. Teff grain structure, anatomy, chemical
composition, and physicochemical properties of the major
chemical components are assessed and described. Teff milling,
processing of injera (the staple food of the majority of Ethio-
pians, a fermented, pancake-like, soft, spongy, sour, circular
flatbread), and other food products made from teff are
described. Lastly, teff potential, grain production limitations,
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00018-8
and recommendations for further research to improve the teff
crop and grain utilization are suggested.
Teff Crop Description and Cultivation
Description
Teff is a C4 self-pollinated tetraploid cereal plant with a chro-
mosome number of 2n 4x 40. Teff is also an allotetraploid
plant. The teff plant and panicles of some varieties are shown
in Figure 1. Teffs root system is fibrous and most stems are
erect, while others are bending or elbowing types. It has a
panicle type of inflorescence showing different forms, from
loose to compact. Its spikelets have 212 florets. Each floret
has a lemma, a palea, three stamens, mostly two ovaries (in
some exceptional cases three), and feathery stigmas. In most
varieties, the plant height is 50120 cm. A single teff plant is
known to produce from 9000 to 90 000 grains depending
upon the variety and production conditions.
Name and Origin
Teff (Eragrostis tef (Zucc.) Trotter) belongs to the family of
Poaceae, subfamily Eragrostoideae, tribe Eragrosteae, and
genus Eragrostis. About 350 species are known in the genus
Eragrostis of which teff is the only cultivated species. Chloridoi-
deae is used synonymously for Eragrostoideae of teff. Vernac-
ular names in different parts of the world are as follows:
Tahf: Arabic
Tef, teff, and Williams love grass: English
Xaafi/tafi/taafi: Oromo (O)/Afar/Sodo; tafe-e: Had; and tef,
teff, and taff: Amarinya (A) and Tigrinya (T) (Ethiopian
languages)
Mil ethiopien: French
Chimanganga, ndzungula (Ch), and chidzanjala (Lo): Malawi
Teff is indigenous to Ethiopia. Ethiopia is also considered the
leading world center for teff genetic diversity. As of 2013, the
Ethiopian Institute of Biodiversity had conserved a total of
5169 accessions and 10 000 teff genotypes for varietal improve-
ment study and to reduce loss of genetic diversity. The exact
details on teff domestication are unclear. However, teff is
believed to have been first domesticated by the pre-Semitic
inhabitants of Ethiopia between 4000 and 1000 BCE and is
assumed to have originated in northeastern Africa. Even
though teff is an allotetraploid plant, to date, its diploid puta-
tive ancestors are not exactly established. But, based on the
morphological data and cytological evidence, the following
species have been suggested as the ancestors and contributors
to teff origin:
Eragrostis aethiopica, E. atrovirens, E. longifolia, E. macilenta,
E. pilosa, and E. pseudo tef as ancestor species of teff
209
210 THE CEREAL GRAINS | Teff: Overview

(a) (c)

(b) (d)

Figure 1 Typical teff field plant and panicles of some teff varieties: (a) teff field of DZ-01-196, (b) panicles of DZ-01-196 (compact), (c) panicles
of DZ-Cr-37 (less compact), and (d) panicles of DZ-01-99 (loose). Reproduced with permission from Tefera, H., Ayele, M., Assefa, K., 1995. Improved
varieties of tef [Eragrostis tef (Zucc.) Trotter] release of 19701995. Research Bulletin 1. Debre Zeit Agricultural Research Center, Ethiopia.

E. aethiopica, E. bicolor, E. cilianensis, E. curvula, E. pilosa, and
E. mexicana as contributor species to the origin of teff
E. aethiopica, E. barrelieri, E. bicolor, E. cilianensis, E. hetero-
mera, E. mexicana, E. minor, E. papposa, and E. pilosa as very
closely related species to teff
E. aethiopica 2; E. barrelieri 6 ; E. cilianensis 2 , 4 , and
6; E. mexicana 6 ; E. minor 2  and 4 ; and E. pilosa 2 
based on cytological evidence as closely related species
to teff
An attempted interspecific cross between teff and some wild
Eragrostis species (E. curvula, E. cilianensis (4 ), E. pilosa (4),
and E. minor) was not successful. However, it was recently
reported that E. tef and E. pilosa can be crossed with fertile
offspring, suggesting that E. pilosa or an ancestor closely related
to E. pilosa is the most probable putative ancestor of teff.
Cultivation and Production
Teff is one of the major cereals in Ethiopia with annual grain
production estimated to be ca. 3.5  106 t (ca. 21% of the
cereal grain production). In Idaho, the United States, some
teff grain is produced for the health-food market and for injera

making. In South Africa, teff is widely grown as a fodder crop
during the summer-rainfall season. However, the production
of a combination of grain and fodder varieties has been
introduced only recently. In Australia, India, Kenya, and the
United States, it is cultivated as a forage crop. Teff has been
started to be cultivated in the Netherlands. Teff can adapt to a
wide range of environments, that is, moisture stress, high
rainfall, different soil types, and a wide range of altitudes
from near sea level to over 3000 m. However, the best condi-
tions are 18002100 m above sea level, a temperature range
of 1027  C during the cultivation period, an annual rainfall
of 750850 mm, and rainfall of 450550 mm during the
growing season. Teff is known to have fewer disease and
pest problems in the field as compared with maize, sorghum,
wheat, and barley. However, the productivity is low. Lack of
high-yielding cultivars, lodging, weeds, waterlogging, low
moisture, low-fertility conditions, loss of grains on harvest-
ing, threshing, and cleaning are major factors that contribute
to the low grain yield. The yields of improved varieties under
improved technologies (fertilizer, weed control, appropriate
harvesting, and grain threshing) on farms are in the range of
1.42.7 t ha1, and for some under research-managed farms,
the yield is as high as 3.6 t ha1. Reduction of seeding rate
from traditionally recommended 2530 kg ha1 to
2.5 kg ha1 had recently demonstrated a substantial grain
yield increase as compared with previous grain yield
recorded. Because of adaption of some of the improved tech-
nologies by teff farmers, the average national yield in Ethiopia
has increased annually from about 0.84 t ha1 in 2003 to
about 1.26 t ha1 in 2011.
Harvesting and Storage
Teff grain is harvested when the vegetative and reproductive
parts (pedicel, lemma, palea, and glumes) turn yellow or
straw color (4560 days for very early maturing, 60120
days for early maturing, and 120160 days for late maturing).
If harvested late, the grain loss will be significant due to
shattering and the natural grain color can also fade. However,
if the grain is harvested early, it may become vitreous or
translucent. In Ethiopia, traditional harvesting is done man-
ually using a sickle (Figure 2(a)). The harvested panicles are
gathered in batches, either on the day harvested or after 13
days, and temporarily stacked (in a conical shape, with the
panicles toward the center) (Figure 2(b)) outdoors, under the
shade, till it is ready for threshing. Such storage duration will
help the teff grain to undergo the ripening process and dor-
mancy breaking stages.
At present, teff grain pre- and postharvest losses are high
(2530%), largely incurred by lodging, shattering on harvest
and harvest handling, threshing, and cleaning by traditional
practices. Threshing is usually done by oxen feet trampling or
manually by beating with sticks after spreading the dried pan-
icles on a dry circular threshing floor. The threshing floor is
prepared by smearing with cow dung, cement, or other suitable
materials. The grain is traditionally winnowed by wafting in
the open air with the help of a rectangular flat piece of dried
leather called an afarsa or hafarsaa (O) (ca. 0.4 m width -
 0.8 m length). Threshers or combine harvesters can be also
used. However, grain loss is large because teff grain is very
THE CEREAL GRAINS | Teff: Overview 211
small and of light mass and can be easily blown away with
the chaff. Use of mechanical threshers has recently shown an
improvement in grain yield and quality. Teff productivity may
be improved in the future by adaption of this technology. In
Ethiopia, the grain is traditionally stored in gotera (A, T) or
gotaara (O) (small hut-like stores) or pots or sacks. In compar-
ison with other common cereals, teff grain is less prone to
attacks by weevils and other storage pests. Thus, it can be safely
stored under traditional storage conditions.
Teff Grain Marketing
In local markets of Ethiopia, several small-scale grain traders
distribute teff grain from major growing areas to the urban
consumers and to regions of shortage in teff grain production.
At Addis Ababa, teff grain is marketed in a place called Ehil
berenda (Markato). Even though teff grain fetches premium
price in the global cereal grain markets, at present, interna-
tional teff grain export from Ethiopia is not practiced because
of limited production. But when permitted, wherever the
market is, companies (government-owned, joint ventures,
farmers union associations, and private firms), such as Ethio-
pian Grain Trading Enterprise and Oromo Development Asso-
ciation, which export oil and pulses, have the potential to
export teff grain as well.
Physicochemical Properties of Teff Grain
Morphology
Teff grain is hull-less (naked) and comes in a range of colors
from milky white to almost dark brown. The most common
colors are white, creamy white, light brown, and dark brown
(Figure 3). The grain is oval-shaped with size 0.91.7 mm
(length) and 0.71.0 mm (diameter). The individual grain
mass is generally 2 mg,  0.60.8% of the wheat grain
mass. The thousand kernel weights and hectoliter weight of
teff grain are in the range 0.190.42 g and 8587 kg hl1,
respectively.
Anatomy
Pericarp
The outer pericarp is thin and membranous and is equivalent
to the beeswing bran of wheat. In the pericarp, beneath the
cuticle toward the nucellar epidermis, teff grain is known to
bear slime layer rich in pectins, which form uniform layers on
the inner surface of the cell wall. The ability of this slime layer
to absorb and maintain moisture around the grain is impli-
cated as a contributor to teff adaptive features to moisture
stress. In the inner surface of the pericarp, the mesocarp and
endocarp are fused and appear as a single layer. In this fused
layer, some starch granules are observed, as in the pericarp of
sorghum grain.
Seed coat (testa)
Next to the endocarp is the testa, which is adjacent to the
aleurone layer. In some teff varieties, the testa is reported to
contain tannins and is thus presumed to be thick. However, in
212 THE CEREAL GRAINS | Teff: Overview
Figure 2 (a) Harvesting of teff plant by the traditional method, (b) stacked
teff panicles after harvest, and (c) teff grain after cleaning by winnowing and
wafting, where TF threshing floor top layer smeared with cow dung and
dried, GS grass straw made of traditional bucket, and IM impurities.
the varieties that the author analyzed, including brown varie-
ties, there is no significant tannin content found.
Aleurone layer
The aleurone layer is one cell thick and is rich in protein and
lipid bodies.
Germ (embryo)
Like in other small-grain cereals, the germ occupies a relatively
large proportion of the grain and is rich in protein and lipids.
Endosperm
The endosperm is the largest component of the grain and
consists of outer and inner layers. The outer layer is vitreous
and contains most of the protein reserves of the endosperm and
a few starch granules. The inner layer is mealy consisting mainly
of thin-walled cells containing mostly starch granules with a
few protein bodies. Teff has compound-type starch granules
(Figure 4(a)), representing the contents of one amyloplast-
like rice, oats, Amaranthus, and quinoa starches. On milling,
individual starch granules are released along with small groups
of protein bodies. The protein bodies are individual entities in
nature and spherical in shape (Figure 4(a)), and unlike those of
wheat, they do not coalesce to form a matrix.
Teff Grain Chemical Composition
The proximate chemical composition of teff grain is shown in
Table 1.
Carbohydrate
Carbohydrate content of teff grain is  73%, of which virtually
all is starch. The teff starch properties are given in Table 2.
Individual starch granules are very small (26 mm in diameter)
(Figures 4(b)4(f)), similar in size to rice starch granules, but
larger than Amaranthus and quinoa starch granules. The shape
is polygonal, smooth with no surface pores (Figure 4(f)). A few
granules are essentially cubic, and at high magnification, some
appear tortoise shell-shaped (Figure 4(e)). The composition of
teff starch granules is similar to other normal native cereal
starches, with 2530% amylose. Gelatinization temperature is
high, similar to other tropical cereal starches. The x-ray diffrac-
tion pattern of teff starch granule is A-type with crystallinity of
 37% similar to rice starch granule. Pasting temperature is
similar to that of maize starch, but cooking time for peak
viscosity is longer. Peak, breakdown, and setback viscosities
are lower than those of maize starch. The paste clarity of
teff starch is opaque. The gel texture is short and smooth.
a-Amylase degradation of teff starch granules is by surface
erosion and endocorrosion in nature.
Because teff starch granules are very small, smooth, and of
uniform size, they offer good functionality as a fat substitute
and a flavor and aroma carrier, similar to other small-granule
starches. Teff starch has good resistance to shear breakdown,
and thus, it may find good application in high-shear processed
foods. Also, because of its slow retrogradation tendency, it
 THE CEREAL GRAINS | Teff: Overview 213

Figure 3 Grains of different teff varieties: (a) DZ-01-196 (white), (b) DZ-Cr-37 (creamy white), (c) DZ-01-99 (light brown), and (d) South African brown
(dark brown).

Table 1 The proximate composition (db) of teff grain, Osborne protein fractions, and food energy

Biochemical class Compound Range (%) Typical value (%)

Protein (%) (N  6.25) 9.413.3 11.0

Carbohydrate (%) 73.0 73.0
Crude fiber (%) 2.03.5 3.0
Fat (%) 2.03.1 2.5
Ash (%) 2.73.0 2.8
Osborne protein fractions (% protein recov.) Albumins 2439 36
Globulins 734 18
Prolamins 315a, 3843b
Glutelins 2842 40
Food energy (kJ per 100 g) 1406 1406
a
Extracted with 60% ethanol.
b
Extracted with tert-butanol containing 0.05% 1,4 dithiothreitol.
Source: Bultosa, G., Taylor, J.R.N., 2004. Teff. In: Wrigley, C., Corke, H., Walker, C.E. (Eds.), Encyclopedia of Grain Science, vol. 3. Elsevier, Oxford, UK, pp. 281290; Ketema, S.,
1997. Tef Eragrostis tef (Zucc.) Trotter. Promoting the Conservation and Use of Underutilized and Neglected Crops, Institute of Plant Genetics and Crop Plant Research, Gatersleben/
International Plant Genetic Resources Institute, Rome, Italy, Booklet 12; Assefa, K., Chanyalew, S., Tadele, Z. (Eds.), 2013. Achievements and Prospects of Tef Improvement:
Proceedings of the Second International Workshop, 79 November, 2011, Debre Zeit, Ethiopia. Ethiopian Institute of Agricultural Research/Institute of Plant Sciences, University of
Bern, Addis Ababa, Ethiopia/Bern, Switzerland; NRC (National Research Council of the USA). 1996. Tef. In: Lost Crops of Africa, vol. 1, Grains, pp. 214338. Washington, DC:
National Academy Press.; Hundera, F., 1998. Variations of morphoagronomic characters and grain chemical composition of released varieties of tef [Eragrostis tef (Zucc.) Trotter]. J.
Genet. Breed. 52, 307311.
214 THE CEREAL GRAINS | Teff: Overview

cw

sg pg
sg

sg
pb

pb pb
cb

(a) (d)

pg
tr

cb

(b) (e)

f cb nsp

pg

(c) (f)

Figure 4 Compound (a) and individual (bf) starch granules from different teff varieties: (a, b) South African brown, (c) DZ-01-1681, and (df)
DZ-01-196 (sg starch granules, pg polygonal, cb cubic, Tr tortoise-shell, pb protein, f fiber, and nsp no surface pores). Reproduced from
Bultosa, G., 2003. PhysicoChemical and Functional Properties of Grain tef [Eragrostis tef (Zucc.) Trotter] Starch. PhD thesis, University of
Pretoria, Pretoria, South Africa.

could have attractive applications where starch staling is pre-
ferred to be reduced (i.e., in baked and in refrigerated foods).
Fiber
The fiber content of teff grain (Table 1) is apparently higher
than most other common cereals, because the grain is very
small and the bran is proportionally large.
Protein and amino acids
Typical teff grain protein content (N  6.25) is  11%, with a
normal range of 913%. Thus, the protein content of teff
grain is similar to other common cereals. The major amino
acids are glutamic acid, alanine, proline, aspartic acid, leu-
cine, and valine (Table 3). Methionine, phenylalanine, and
histidine are slightly higher than in most other cereals, but

Table 2 Physicochemical properties of teff grain starch
Property
Individual granule diameter (mm) and shape
Amylose (%) (db)
Amylose weight-average molar mass, g mol1
Amylopectin weight-average molar mass, g mol1
Ash (%) (db)
Protein (%) (N  6.25) (db)
Lipids (db): total (mg g1)
Internal lipids (mg g1)
Phosphorus (db) (mg g1)
Kofler gelatinization temp. range ( C): ToTpTc
DSC gelatinization endotherms range: To, Tp, and Tc in

C and DH in J g1, respectively
Pasting properties: Ti ( C), PV (RVU), BV (RVU), Rst
(RVU min1), SBV (RVU)
Peak viscosity
Resistance to shear
Retrogradation tendency
Crystallinity (%)
x-Ray diffraction: d-value (A) and intensity (%)
Paste clarity
Gel texture
In vitro digestibility with porcine pancreatic a-amylase
Acid hydrolysis
Source: Bultosa, G., 2003. PhysicoChemical and Functional Properties of Grain tef [Eragrostis tef (Zucc.) Trotter] Starch. PhD thesis, University of Pretoria, Pretoria, South Africa;
Bultosa, G., Hamaker, B.R., BeMiller, J.N., 2008. An SEC-MALLS study of molecular features of water-soluble amylopectin and amylose of tef [Eragrostis tef (Zucc.) Trotter] starches.
Starch/Starke 60, 822.
serine and glycine are lower. Lysine and arginine are essen-
tially higher in teff than in most other cereals, except rice and
oats. The balance among essential amino acids is similar to
the whole edible portion of egg protein, except for its lower
lysine content. The overall amino acid profile of teff can be
regarded as well balanced.
Osborne protein fractions of teff grain are shown in Table 1.
Glutelins, albumins, and globulins are major fractions. The teff
prolamin fractions are reported variable (low and high)
depending on solvent type used in the extraction. The major
prolamins of teff are somewhat similar to the a-prolamins of
maize, sorghum, and Coix. Teff is different from other cereals
in having higher albumins and globulins. Teff protein is essen-
tially free of the type of gluten found in wheat. Because of this,
teff grain foods are today increasingly became important for
consumers allergic to wheat gluten (e.g., for celiac patients). As
the main protein fractions (albumins and globulins) are the
most digestible types, teff protein digestibility is also presumed
high. Amino acid compositions of the various Osborne protein
fractions are shown in Table 3.
Ash and minerals
The ash content in teff grain (Table 1) is apparently higher
than in wheat, rye, maize, barley, oats, rice, and millets, in part
THE CEREAL GRAINS | Teff: Overview 215
Mean and standard deviation for five teff
varieties Remark
26 (majority are 35), polygonal shape, Individuals are from compound granules
smooth surface
28.4  2.8 Concanavalin A method
28.2  0.8 Iodine-binding method
(1.53.0)  106
(10.116.5)  107
0.16  0.04
0.19  0.03
8.9  0.7 24% HCl hydrolysis followed n-hexane and
in situ generated ethyl formate extract
7.8  0.4 Hot (90  C) water-saturated butanol extract
0.65  0.08
68.074.080.0 To is onset, Tp is peak, and Tc is conclusion
gelatinization temperatures
63.865.4, 70.271.3, and 81.381.5 and DH is gelatinization enthalpy
2.287.22
74.0  1.1, 269  13, 79  17, 8.4  1.8, Where Ti, PV, BV, Rst, and SBV are pasting
101  11 temperature, peak viscosity, breakdown
viscosity, rate of shear thinning, and
setback viscosity, respectively
Mediumlow
Mediumhigh
Mediumlow
37
5.85 (83.8), 5.16 (97.0), 4.89 (99.4), 4.41 A-type starch crystalline polymorph
(36.4), and 3.84 (80.2)
Opaque
Short
Surface erosion and endocorrosion type
Gradual surface degradation (etching)
because teff grain bran is proportionally large. In particular,
calcium, copper, iron, and zinc (Table 4) contents are higher
compared with those in barley, wheat, and sorghum. The iron
content of traditionally harvested teff grain is especially high
( 15.7 mg per 100 g and in some reports as high as 27 mg per
100 g), in part because of grain contamination with the soil
during harvest. However, when cleaned (with water and/or
dilute acid), the level ( 5.7 mg per 100 g) is similar to other
cereals. Most teff foods such as injera are fermented. Fermen-
tation for injera processing is known to destroy phytic acid to
the extent of 90%, and such destruction is known to contrib-
ute to high iron bioavailability in diets where fermented teff
foods are the staples. Because of these two factors, iron-
deficiency disease anemia is rare among regular teff injera
consumers in Ethiopia.
Fat and fatty acids
Teff grain fat (Table 1) is lower than, for example, in maize and
oats. Thus, teff is different from other small-grain cereals in
having low fat even though the germ is large. As in most other
cereal grains, palmitic, oleic, and linoleic acids are the major
fatty acids (Table 5). Linolenic acid in teff is higher than in
maize, sorghum, and wheat.
216 THE CEREAL GRAINS | Teff: Overview

Table 3 Amino acid composition of the whole teff grain protein and of the Osborne protein fractions (g per 100 g protein)

Asx is Asp Asn and Glx is Glu Gln

Whole grain Albumin Globulin Glutelin Prolamin

Amino acid Range Mean Range Mean Range Mean Range Mean Range Mean
Asx 5.87.2 6.4 10.714.4 12.7 10.314.1 12.9 12.714.4 13.7 3.620.7 9.8
Thr 2.44.4 3.6 3.34.5 4.2 0.04.2 2.6 2.04.2 3.3 2.84.1 3.5
Ser 2.85.6 4.1 3.13.9 3.6 0.12.4 1.3 0.82.2 1.6 3.44.3 3.8
Glx 18.724.9 21.8 3.910.5 9.0 1.59.8 6.9 6.59.5 8.7 4.826.3 20.6
Pro 5.111.4 8.2 6.711.7 8.3 8.016.6 10.6 8.011.0 8.8 6.813.5 10.0
Gly 1.74.1 3.1 4.18.4 7.2 1.85.3 3.9 3.55.4 4.5 4.36.0 4.9
Ala 5.514.7 10.1 13.121.7 15.4 13.323.7 17.2 13.517.9 14.6 10.519.3 12.7
Cys 0.52.5 1.8 0.40.8 0.6 0.42.1 0.9 0.41.3 0.8 0.01.3 1.3
Val 4.19.9 5.9 8.09.6 8.6 9.211.4 10.5 9.210.9 10.0 7.39.2 7.8
Met 2.04.6 3.3 0.50.8 0.7 0.41.2 0.6 0.40.7 0.6 0.72.8 1.8
Ile 3.25.4 4.0 3.44.7 4.2 3.45.1 4.5 3.95.1 4.5 2.54.6 4.0
Leu 6.09.7 8.1 6.16.5 6.3 5.47.4 6.5 6.67.4 7.0 5.17.3 6.5
Tyr 1.74.0 3.0 0.92.5 1.3 0.91.6 1.3 0.81.7 1.3 1.34.8 3.4
Phe 2.75.9 5.0 2.33.5 2.9 2.13.9 3.0 2.63.9 3.2 2.56.6 5.1
His 2.13.7 2.8 2.84.2 3.4 3.65.6 4.6 4.55.1 4.8 0.73.7 1.9
Lys 1.44.0 3.0 3.76.6 5.6 1.44.2 3.0 2.53.9 3.2 0.32.7 1.3
Arg 2.96.2 4.5 5.87.3 6.4 8.413.7 10.3 8.910.7 9.5 1.25.3 2.8
Trp 1.3 1.3

Source: Bultosa, G., Taylor, J.R.N., 2004. Teff. In: Wrigley, C., Corke, H., Walker, C.E. (Eds.), Encyclopedia of Grain Science, vol. 3. Elsevier, Oxford, UK, pp. 281290; Ketema, S.,
1997. Tef Eragrostis tef (Zucc.) Trotter. Promoting the Conservation and Use of Underutilized and Neglected Crops, Institute of Plant Genetics and Crop Plant Research, Gatersleben/
International Plant Genetic Resources Institute, Rome, Italy, Booklet 12; Assefa, K., Chanyalew, S., Tadele, Z. (Eds.), 2013. Achievements and Prospects of Tef Improvement:
Proceedings of the Second International Workshop, 79 November, 2011, Debre Zeit, Ethiopia. Ethiopian Institute of Agricultural Research/Institute of Plant Sciences, University of
Bern, Addis Ababa, Ethiopia/Bern, Switzerland; Hundera, F., 1998. Variations of morphoagronomic characters and grain chemical composition of released varieties of tef [Eragrostis
tef (Zucc.) Trotter]. J. Genet. Breed. 52, 307311; Bekele, E., 1995. Essential and non-essential amino acids in a free state and in the major protein fractions of teff seeds. SINET:
Ethiop. J. Sci. 18, 7999.

Table 4 Microelement composition of teff grain (db) Vitamins

In teff (Table 6), thiamin content is typically lower, when
Microelements (mg per 100 g or compared with that in wheat, rye, barley, oats, rice, maize,
mg per 100 g) Range Typical value millet, and sorghum. Though riboflavin content is considered
to be high, it is nevertheless lower than in rye, barley, and oats.
Calcium (mg) 104223 165.2
Niacin levels are similar to those in maize.
Chloride (mg) 13 13.0
Chromium (mg) 250 250.0
Copper (mg) 0.75.3 2.6
Iron (mg) 4.719.6 5.7a, 15.7b
Magnesium (mg) 138190 169.8 Processing and Usage of Teff Grain
Manganese (mg) 1.66.4 3.8
Phosphorus (mg) 378480 425.4 Cleaning
Potassium (mg) 330570 380.0 Normally, the grain is cleaned by manual sifting and sieving.
Sodium (mg) 11.847.0 15.9
Zinc (mg) 2.06.7 4.8
a
Mean of iron from cleaned, acid- and/or water-washed samples.
b
Milling
Mean of iron from uncleaned samples.
Source: Ketema, S., 1997. Tef Eragrostis tef (Zucc.) Trotter. Promoting the Conservation The cleaned grain is usually dry-milled to obtain whole flour.
and Use of Underutilized and Neglected Crops, Institute of Plant Genetics and Crop Traditionally in Ethiopia, this was done by wafcho (A) (T) or
Plant Research, Gatersleben/International Plant Genetic Resources Institute, Rome, Italy, wafcoo (O) (top and bottom hard stones). Today, milling
Booklet 12; Assefa, K., Chanyalew, S., Tadele, Z. (Eds.), 2013. Achievements and by hand, using hard stone, has been replaced by grist mills
Prospects of Tef Improvement: Proceedings of the Second International Workshop, 79
run by electric power and, where electric power is not available,
November, 2011, Debre Zeit, Ethiopia. Ethiopian Institute of Agricultural Research/
by diesel engine or water power. The grist mill is made up of
Institute of Plant Sciences, University of Bern, Addis Ababa, Ethiopia/Bern, Switzerland;
Bultosa, G., 2003. PhysicoChemical and Functional Properties of Grain tef [Eragrostis two abrasive hard-disk stones. During operation, one stone is
tef (Zucc.) Trotter] Starch. PhD thesis, University of Pretoria, Pretoria, South Africa; NRC stationary, while the other is rotating. The grain, fed into the
(National Research Council of the USA). 1996. Tef. In: Lost Crops of Africa, vol. 1, center (eye) of the upper stone, is fragmented and ground
Grains, pp. 214338. Washington, DC: National Academy Press.; Umeta, M., West, between the two stones, and flour is issued at the periphery.
C.E., Fufa, H., 2005. Content of zinc, iron, calcium and their absorption inhibitors in At present, wet milling of teff grain for chemical component
foods commonly consumed in Ethiopia. J. Food Comp. Anal. 18, 803817. extraction like starch is not carried out.
 THE CEREAL GRAINS | Teff: Overview 217

Table 5 Fatty acid composition of teff grain fat plasticity losses after a day. The rice flour injera-making ability
is similar to teff probably because of somewhat similar starches
Fatty acids (%) Range Mean in both grains. The hydrophobic and less polymerized nature
Palmitic (C16:0) 14.016.4 15.9 of teff grain protein prolamins is in part implicated as contrib-
Palmitoleic (C16:1) 0.10.6 0.3 utor to the making of superior semileavened flatbread injera
Stearic (C18:0) 3.03.7 3.3 when compared with sorghum injera. Teff grain flour is also
Oleic (C18:1) 23.324.9 24.0 used to make sweet unleavened bread called kitta (A), bixxille
Linoleic (C18:2) 41.346.5 44.2 (O), and daguwalo (T). Kitta can be consumed as bread, or it
Linolenic (C18:3) 6.99.9 7.9 can be used as an adjunct in traditional opaque beer (tella (A)
Arachidic (C20:0) 0.60.9 0.7 or farsoo (O)) or local spirit (katikalla (A) or araqii (O)). Por-
Arachidonic (C20:1) 0.51.2 0.8 ridge (genefo (A) or marqaa (O)) can also be made from teff
Behenic (C22:0) 0.31.1 0.5
flour. Thin, fermented teff flour batter is used to prepare soup
Erucic (C22:1) 0.00.9 0.4
(muk, (A)). Unfermented teff flour dough is also used in the
Source: Hundera, F., 1998. Variations of morphoagronomic characters and grain preparation of traditional snacks (dabo kolo (A) or hunkuroo
chemical composition of released varieties of tef [Eragrostis tef (Zucc.) Trotter]. J. (O)), where the dough is rolled into small balls and then
Genet. Breed. 52, 307311. roasted on a hot griddle. In the United States, teff has been
promoted as a thickener for soups, stews, and gravies probably
because teff flour paste gives the product a short and stiff
texture. Teff grain flour imparts a slight molasses-like sweetness
Table 6 Vitamin and antinutrient composition of teff grain (db)
to food products, making its inclusion in porridges, pancakes,
Component Typical value biscuits, cookies, cakes, stir-fry dishes, casseroles, soups, stews,
and puddings desirable. Processing of teff grain in Ethiopia has
Vitamins been limited to the household level. To date, technologies for
Vitamin A (mg per 100 g as RE) 8 large-scale commercial processing of teff grain, for the prepa-
Thiamine (mg per 100 g) 0.3 ration of foods like injera, are not well advanced. However,
Riboflavin (mg per 100 g) 0.2
apart from traditional usage, recent reports indicate that teff
Niacin (mg per 100 g) 2.5
grains, along with soybean, chickpea, and other grains, are
Vitamin C (mg per 100 g) 88
Antinutrients being used in the baby food industry. Teff grain flours are
Phytate (mg per 100 g) 707 used to improve dietary fiber, starch, protein, and mineral
Trypsin inhibitor activity (TIU per gram) 5584 content supply in gluten-free food products and in wheat
bread to improve the iron and antioxidant contents of the
RE is retinol equivalent; TIU is trypsin inhibitor unit. bread.
Source: Assefa, K., Chanyalew, S., Tadele, Z. (Eds.), 2013. Achievements and Prospects
of Tef Improvement: Proceedings of the Second International Workshop, 79
November, 2011, Debre Zeit, Ethiopia. Ethiopian Institute of Agricultural Research/ Teff Grain Injera Making
Institute of Plant Sciences, University of Bern, Addis Ababa, Ethiopia/Bern, Switzerland;
Bultosa, G., 2003. PhysicoChemical and Functional Properties of Grain tef [Eragrostis In Ethiopia, injera is regarded as the national staple food.
tef (Zucc.) Trotter] Starch. PhD thesis, University of Pretoria, Pretoria, South Africa; NRC Flowcharts of the injera-making process and injera-baking pro-
(National Research Council of the USA). 1996. Tef. In: Lost Crops of Africa, vol. 1, cess are shown in Figures 5 and 6, respectively. The process
Grains, pp. 214338. Washington, DC: National Academy Press. involves fermentation and then baking of the batter.

Usage
Food made from teff grain is a staple diet for many Ethiopians.
Teff is considered to have a better food value than major grains,
namely, wheat, barley, sorghum, and maize, as it is normally
used as a whole grain, that is, the germ and bran are consumed
along with the endosperm. Teff flour is used primarily for
making of injera (A), caabitaa or budeena (O), and tayeta (T).
Teff injera is regarded to have pre- and probiotic potentials due
to dietary fiber from whole grain and fermentation,
respectively. Although the lactic acid bacteria (LAB) and yeast
involved in the fermentation of injera are killed during baking,
their dead cells and metabolic products are consumed as part
of the diet, and these were implicated as promoters of gut
health. In injera-making features, teff grain flours are superior
to any other cereal grain flours used. The Ethiopian diaspora is
using injera from rice or blend of rice and wheat flour,
although the injera suffers from staling, resilience, and
Fermentation
Flour is mixed with water and the dough is kneaded, usually by
hand. Fermentation for injera making involves two phases that
can last a total of 2472 h. The first phase starts spontaneously
when the flour is wetted, due to contaminating microorgan-
isms. Or it can be initiated by addition of irsho (A) (T) or
raacitii (O) (yellowish liquid saved from the previous batch
fermentation). The initial 1824 h is notable for vigorous gas
evolution and maximum dough expansion. At about 3033 h,
an acidic yellowish liquid appears on the dough surface. This
phase is characterized by a decrease in gas evolution up to 31 h,
an increase in liquid volume up to 48 h, and a decrease in pH
to below 5.8.
The first phase of fermentation results in a liquid/solid
separation after  24 h. The layer of liquid is then removed.
About 10% of the fermenting dough is mixed with water (1:3
ratio) and boiled (25 min), and as a result of starch gelatini-
zation, a dough binder, called absit, is formed. The absit is
cooled and added to the fermentation vat signaling the second
phase of fermentation. The second phase (0.52 h) is
218 THE CEREAL GRAINS | Teff: Overview

Teff grain

Cleaning

Millings

Irsho (starter, yellowish liquid saved

Water (ca 6 L) Teff flour (ca. 3 kg) from previous fermentation)
(ca. 480 g)

Fermented paste separation

(ca. 3033 h) Supernatant (irsho)

First phase fermentation

Sediment
(2472 h) Mixing of 10% sediment with water
(90%)
(1:3) and cooking for 25 min

Air cooling to not less than 46C

Absit (a dough binder)

Second phase fermentation
(0.52 h)

Thinning of fermented batter Water

Pouring of the batter on hot griddle

and covering with griddle lid

Heat and steam baking (23 min)

Injera (18 total each weigh ca. 450 g)

Figure 5 Flowchart of teff injera-making process.

characterized by a short duration of dough expansion and gas
formation.
During the first phase of fermentation, the yellowish liquid
that is removed contains water-soluble nutrients (amino acids,
sugars, and minerals) and a large number of microorganisms
involved in the fermentation. This has negative nutritional
consequences. Thus, injera baked from a batter at  31 h of
fermentation without discarding the liquid is recommended as
being more nutritious.
A complex group of microorganisms is known to be
involved in teff fermentation. A total of 107 LAB (some hetero-
fermentative and some homofermentative) and 68 yeast
strains are reported in the teff fermentation. Bacteria belonging
to the Enterobacteriaceae family are thought to initiate the
fermentation. During the first 18 h of fermentation, the activ-
ities of these bacteria reduce the dough pH to 5.8. A group of
LAB of which major ones are Leuconostoc mesenteroides, Strepto-
coccus faecalis, Pediococcus cerevisiae, P. pentosaceus, Lactobacillus
brevis, L. plantarum, and L. fermentum are involved at the later
phase of fermentation (1872 h) in reducing the dough pH
from 5.8 to 3.8. During the later phase of fermentation (22
24 h), yeasts of two genera Saccharomyces and Torulopsis are
reported to be involved. In the later phase ( 48 h), yeasts
belonging to the genera Candida and Pichia are the dominant
types isolated from the yellowish liquid removed from the
dough. In addition to amylases present in the grain, the

DZ-01-787
Figure 6 Teff injera baking from fermented batter on electrically heated baking griddle (a). Portion of baked injera from white (b and c) and brown (d)
teff grains.
bacteria species Bacillus sp. A-001 involved in the fermentation
has been characterized as one of the amylase-producing bacte-
ria involved in partially attacking the starch granules.
In the traditional teff fermentation for injera making, com-
mercial yeast is not added externally. The source of the yeast is
either from irsho or from the endogenous microflora of teff
grain and its flour that grows in the batter after the flour is
wetted with water. Therefore, the yeast in the fermentation of
teff for injera making can be regarded as symbiotic yeast.
Baking
Injera is usually baked after 24 h of fermentation. After the
fermentation, the batter is diluted slightly with water and then
poured using circular motion from the outer perimeter toward
the center onto a hot, round, smooth griddle called a metad (A)
or eelee caabitaa (O). It is then covered with a metad lid called
akambalo (A), qadaada eelee caabitaa (O) to prevent steam from
escaping. The griddle is traditionally made from clay. Before
pouring the batter, the metad surface is swabbed with ground
oilseeds, commonly rapeseed or with animal fat in a piece of
cloth. This prevents the injera from sticking to the metad sur-
face. Depending on the batter thickness, heat intensity applied,
and steaming, with the current electrically heated baking grid-
dle, injera can be baked in 23 min.
Based on the duration and nature of the fermentation
involved, three common types of injera are prepared:(1) injera
made from dough that does not contain absit, characterized by
a soft, thin, fine appearance and a sour taste without the eyes
of injera (surface air cells); (2) injera made from partially
fermented paste (1224 h fermentation) called aflegna (A)
(O) or bekuo (T), characterized by a sweet flavor, a pleasant
odor, and a rusty red underside; and (3) injera made from
overfermented paste called komtata injera (A) (T) or qomxoxaa
caabitaa (O) that tastes very sour and is regarded as less
THE CEREAL GRAINS | Teff: Overview 219
DZ-01-196
DZ-01-99
nutritious. The thick batter used for aflegna injera is also used
to prepare a slightly concave, thick flatbread called cumboo (O).
Cumboo is traditionally baked on the preheated surface of a
small-size concave griddle called bedde, which is placed upside
down on a flat larger griddle called elle chumbo. After pouring
the 1224 h fermented thick batter onto the upside down
concave bedde surface and covering with the lid, gentle heat is
applied underneath of the larger griddle, and depending on the
batter size and heat intensity applied, the baking time for
cumboo is between 1 and 3 h.
Prospects, Problems, and Recommendations
Teff can be cultivated under harsh environmental conditions
where most other cereals are less viable. It has relatively few
pest and disease problems in the field. The grain is less attacked
by weevils. The nutrient composition of teff grain indicates that
it has good potential to be used in various diets and as func-
tional foods in health-food markets worldwide. However, the
grain yield of teff is low. The mechanized farming technologies
that are used for the production of other cereal grains can be
problematic for teff because the plant stems are very thin and
short and the grain is very small. Manipulation of teff plant
genes through research on germplasm genetic diversity,
improvement of yield using molecular breeding approach,
and a continuous teff production technology improvement
for maximum grain production are required.
At present, the milling of teff grain is limited to cottage-type
millers. Processing of teff for different foods is usually by
traditional ways and mostly limited to the household level.
Over the last decade, utilization of teff grain as whole grain
supplements and for gluten-free markets is increased. More
research on teff large-scale milling and processing of teff grain
220 THE CEREAL GRAINS | Teff: Overview
for different commercial foods is required to promote world-
wide teff product utilization.
Exercises for Revision
What major features distinguish teff grain (particularly its
starch and protein) from other common tropical cereals
like sorghum, rice, and maize?
Why is the productivity of the teff plants so low compared
with that of maize and rice? Could teff be grown near where
you live?
Why are teff grain products nutritionally and functionally
attractive for health-food markets (e.g., for celiac patients)?
Can you design and develop a novel food product utilizing
teff grain?
How is injera made? Is teff grain essential for injera manufac-
ture? Does teff injera have prebiotic and probiotic effects?
Exercises for Readers to Explore the Topic Further
Value chain efficiency for teff productivity improvement
such as high-yielding and lodge-resistant variety develop-
ments coupled with mechanized or semiautomated pro-
duction technologies that suit very thin, short teff plant
stems and very small grain size of light mass that would
boost teff grain production and minimizes grain posthar-
vest losses.
Development of teff grain-based composite flours for injera
and other teff products.
Efficiency of teff injera processing stages and cottage and
large-scale injera processing and preservation.
Teff grain-based ready to eat foods for health-food markets
such as gluten-free foods for celiac patients.
See also: Beverages from Grains: Fermentation: Foods and
Nonalcoholic Beverages; Food Grains and the Consumer: Cultural
Differences in Processing and Consumption; Grains and Health; Food
Grains and Well-being: Functional Foods: Dietary Fibers,
Prebiotics, Probiotics, and Synbiotics; Functional Foods: Overview;
Food Grains: Intolerance, Allergy and Diseases: Celiac
Disease; The Gluten-Free Diet; Appendix 1: Nutrient-Composition
Tables for Grains and for Grain-Based Products; Grain Composition
and Analysis: The Composition of Food Grains and Grain-Based
Products; Grains Around the World: Grain Production and
Consumption: Africa; The Basics: Grain and Plant Morphology of
Cereals and how characters can be used to identify varieties; The Grain
Crops: An Overview; Grain: Morphology of Internal Structure;
Taxonomic Classification of Grain Species; The Cereal Grains: An
Overview of the Family of Cereal Grains Prominent in World
Agriculture; Millet Minor: Overview; Oats: Overview; Rice: Overview;
The Legumes and Pseudocereals: Amaranth: Overview.
Further Reading
Adebowale AA, Emmambux MN, Beukes M, and Taylor JRN (2011) Fractionation and
characterization of tef proteins. J. Cereal Sci. 30: 17.
Assefa K, Chanyalew S, and Tadele Z (eds.) (2013) Achievements and Prospects of Tef
Improvement: Proceedings of the Second International Workshop, November 79,
2011, Debre Zeit, Ethiopia, Addis Ababa, Ethiopia/Bern, Switzerland: Ethiopian
Institute of Agricultural Research/Institute of Plant Sciences, University of Bern.
Bekele E (1995) Essential and non-essential amino acids in a free state and in the major
protein fractions of teff seeds. SINET: Ethiop. J. Sci. 18: 7999.
Bergamo P, Maurano F, Mazzarella G, Iaquinto G, Vocca I, Rivelli AR, et al. (2011)
Immunological evaluation of the alcohol-soluble protein fraction from gluten-free
grains in relation to celiac disease. Mol. Nutr. Food Res. 55: 12661270.
Bultosa, G., 2003. PhysicoChemical and Functional Properties of Grain tef [Eragrostis
tef (Zucc.) Trotter] Starch. PhD thesis, University of Pretoria, Pretoria, South Africa.
Bultosa G and Taylor JRN (2004) Teff. In: Wrigley C, Corke H, and Walker CE (eds.)
Encyclopedia of Grain Science, vol. 3, pp. 281290. Oxford, UK: Elsevier Ltd.
Bultosa G, Hamaker BR, and BeMiller JN (2008) An SEC-MALLS study of molecular
features of water-soluble amylopectin and amylose of tef [Eragrostis tef (Zucc.)
Trotter] starches. Starch/Starke 60: 822.
Desiye A and Abegaz K (2013) Isolation, characterization and identification of lactic acid
bacteria and yeast involved in fermentation of teff (Eragrostis tef) batter. Adv. Res.
Biol. Sci. 1(3): 3644.
Ebba T (1969) Tef (Eragrostis tef): the cultivation, usage and some of the known
diseases and insect pests. Part I. Debre Zeit Agricultural Experiment Station. Dire
Dawa, Ethiopia: Addis Ababa University, College of Agriculture.
Hundera F (1998) Variations of morpho-agronomic characters and grain chemical
composition of released varieties of tef [Eragrostis tef (Zucc.) Trotter]. J. Genet.
Breed. 52: 307311.
Ketema S (1997) Tef Eragrostis tef (Zucc.) Trotter. Promoting the Conservation and Use
of Underutilized and Neglected Crops. Gatersleben/Rome: Institute of Plant Genetics
and Crop Plant Research/International Plant Genetic Resources Institute Booklet 12.
Kreitschitz A, Tadele Z, and Gola EM (2009) Slime cells on the surface of Eragrostis
seeds maintain a level of moisture around the grain to enhance germination. Seed
Sci. Res. 19: 2735.
NRC (National Research Council of the USA) (1996) Tef. In: Lost Crops of Africa, vol. 1.
Grains, pp. 214338. Washington, DC: National Academy Press.
Tefera H, Ayele M., Assefa, K., 1995. Improved varieties of tef [Eragrostis tef (Zucc.)
Trotter] release of 19701995. Research Bulletin 1. Debre Zeit Agricultural Research
Center, Ethiopia.
Umeta M and Parker ML (1996) Microscopic studies of the major macro-components of
seeds, dough and injera from tef (Eragrostis tef). SINET: Ethiop. J. Sci.
19: 141148.
Umeta M, West CE, and Fufa H (2005) Content of zinc, iron, calcium and their
absorption inhibitors in foods commonly consumed in Ethiopia. J. Food Comp.
Anal. 18: 803817.
Relevant Websites
http://www.ars-grin.gov USDAs GRIN database.
http://mansfeld.ipk-gatersleben.de Mansfelds world database of agricultural and
horticultural crops.
http://www.tef-research.org/-Tef Research project.
 The Oilseeds

Contents
Oilseeds: Overview
Soybean: Overview
Canola: Overview
Cottonseed: Overview
Sunflower: Overview
Overview of the Oilseed Safflower (Carthamus tinctorius L.)
Linseed: Overview

Oilseeds: Overview
RJ Mailer, Australian Oils Research, Lambton, NSW, Australia
2016 Elsevier Ltd. All rights reserved.

Topic Highlights Introduction

The major oilseed crops of the world include soybean,
cottonseed, rapeseed, sunflower, groundnut, sesame seed,
linseed, safflower, and mustard seed.
The global oilseed market is dominated by soybean with a
total of 242 Mt produced in 2012.
Rapeseed is now rated in second position in world produc-
tion of oilseeds with 65 Mt produced in 2012.
Plant-breeding programs are aimed at increasing oil pro-
duction and quality.
World oilseed production is forecast to expand by 20% for
the period 201221.
The use of edible oil for biodiesel is forecast to increase by
76% or 30 Mt by 2021.
Meal is utilized as stock feed, in particular pork and poultry
and, more recently, aquaculture.
Palm oil fruit production in 2012 exceeded 50 Mt, second
only to soybean.
The biggest changes will come in the future from genetic
modification.
Oilseed crops are generally grown for the oil in their seeds and
vary considerably in oil content, quality, and composition
factors that rely heavily on the crop species or cultivar and
upon the environmental conditions in which the crop is
grown. The seed meal left after the oil has been extracted can
provide a high-protein product for use in either animal or
human food. In some crops, the seed meal or some other
factors are more important than the oil, and the oil is a by-
product. For example, soybean is primarily grown for its high-
protein meal and cotton for its fiber. Consequently, in
oil-producing crops, plant-breeding programs are aimed at
increasing both oil production and quality with the additional
aim of increasing meal quality. In some oil crops, antinutri-
tional components are present (such as gossypol in cottonseed
and glucosinolates in rapeseed); they also need to be consid-
ered and may limit the use of the meal.
Oilseeds and Their Uses

The major component of oilseed crops is the oil, which, in

Learning Objectives
This article is designed to provide the reader with some
basic information about the major oilseed crops of the
world.
The reader should have a basic understanding of the impor-
tance of each crop and its uses.
Information on production and export are provided to
show which countries are the major producers and
exporters.
many cases, accounts for up to 80% of the crop value. The oil-
free meal provides additional value along with various by-
products, such as lecithin, and a range of extracts from both
meal and oil.
Vegetable oils contain 9598% triacylglycerols (or triglyc-
erides). The remaining fraction consists of phospholipids,
monoacylglycerols and diacylglycerols, and unsaponifiable
components including sterols and tocopherols. The oils consist
of long-chain fatty acids from C14 to C24 in length, and the
proportion of these fatty acids in the oil has a significant
contribution to its nutritional value. In recent years, there has

Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00026-7 221

222 THE OILSEEDS | Oilseeds: Overview
been a trend in human diets toward reduction of saturated fats
such as palmitic acid (C16:0) and an increase in polyunsatu-
rated (C18:2 and C18:3) and monounsaturated (C18:1) fats.
Plant breeders have successfully developed more nutritionally
favorable products from traditional oilseed crops by altering
their fatty acid profiles and other constituents.
There is strong demand for food, feed, and biofuel. According
to the OECD-FAO Agricultural Outlook 201221, world oilseed Rapeseed
production is forecast to expand by 20% for the period, exceed-
ing 529 Mt by 2021. This is a marked slowdown compared with
that of the last decade and is as a result of reduced area expan-
sion, environmental constraints, and competing crops. Vegeta-
ble oil production, however, will increase by about 28% due to
the contribution of palm and coconut oil. By 2021, palm oil is
expected to make up one-third of world vegetable oil production.
World consumption of oils is forecast by FAO to grow by only
2% per year, half the growth rate of the last decade. China and
the European Union are the dominant importers of oilseeds and
their products. Although demand for food use is slowing down
in developed countries as they reach saturation levels, consump-
tion in developing countries is steadily increasing. There has
been and continues to be a significant increase in biodiesel use
in developed and developing countries. The use of edible oil for
biodiesel is forecast to increase by 76% or 30 Mt by 2021, raising
the share of vegetable oil for biodiesel from 12% to 16%. Non-
food uses of vegetable oils in developed countries are estimated
to continue to increase due to developments in the oleochemical
industries with a shift away from petroleum-based products to
environmentally friendly oleochemicals. In addition, there is an
increase in technological developments, particularly with the
genetic modification of oilseeds to create a new range of prod-
ucts. For cooking oils and salad dressings, plant breeders have
selected cultivars with lower levels of polyunsaturated and satu-
rated fats and an increase in monounsaturates for increased
oxidative stability. New developments include oils with
increased stearic acid to provide margarine type fats without
the need for hydrogenation and the subsequent production of
trans-fatty acids. The uses of vegetable oils in paints, lubricants,
cosmetics, and pharmaceuticals have been well documented.
Meal production globally will increase by 23% or about
315 Mt by 2021, much of this as a result of expansion of
rapeseed for biodiesel in the European Union. World con-
sumption of oilseed meal is also expected to increase by 23%,
which compared with the past decade reflects a slowing down
with livestock industries growing at a slower pace. Soybean
meal dominates the market although it is expected that future
meal production from sunflower and rapeseed will increase
relative to soybean. Much of the meal is utilized as stock feed
and in the production of livestock products, in particular pork
and poultry and, more recently, aquaculture. Meal products,
such as protein extracts, are used for edible purposes, and
consumption of whole seed, including roasted soybeans, pea-
nuts, and sesame seed, is common. The meal also has indus-
trial applications in cosmetics, paints, and adhesives.
Oil Analysis
The traditional methods of oilseed analysis made plant breed-
ing and selection a slow and tedious process. The Goldfisch or
Soxhlet methods of oil extraction required several hours to
Table 1 Approximate oil content of seed and protein content
of oil-free meal and the main uses of selected oilseed crops
Oilseed Oil (%) Protein (%) Main use
Soybeans 20 46 Food
Cottonseed 16 37 Fiber
Peanut 41 46 Food
41 34 Oil
Sunflower seed 40 28 Oil
Sesame seed 4060 42 Food
Linseed 40 3234 Oil
Safflower seed 34 23 Oil
Mustard seed 2050 35 Condiment/oil
Source: Adapted from Australian Oilseed Federation Technical and Quality
Standards 2014 and other literature values.
complete and involved the use of hazardous flammable
solvents. In addition, only a few samples could be processed
at a time reducing the capacity of the laboratories to service the
needs of plant-breeding programs. Kjeldahl nitrogen analysis
to determine the nitrogen content of the meal also required the
use of dangerous chemicals and strong acids. More recent
advances in oil content determination include nondestructive
seed analysis, such as the use of nuclear magnetic resonance
spectroscopy. The rapid and accurate nondestructive testing
of seed for oil content allows the breeder to retain the seed of
potentially suitable lines. The introduction of near-infrared
reflectance spectroscopy has further enhanced the ability of
laboratories to determine a wider range of seed components
including oil, protein, moisture, fiber, and numerous others on
relatively small samples in a short time. Consequently, hun-
dreds of seed samples can now be analyzed each day and the
intact seed returned to the breeder for further growth and
evaluation.
New Variations on Old Crops
The major oilseed crops of the world include soybean, cotton-
seed, rapeseed, sunflower, groundnut (or peanut), sesame
seed, linseed (from which the name linolenic acid is derived),
safflower, and mustard seed. Many other crops can be used for
oilseed production including castor beans, grape seed, tobacco
seed, flax, corn oil, tung beans, and okra. The oil and protein
content of the major oilseed crops is shown in Table 1.
From these crops, plant breeders have developed specialty
oil types within the different species. Sunflower oil, for exam-
ple, is traditionally high in linoleic (polyunsaturated) acid and
is promoted as such for its health benefits. With further plant
breeding, there are now several grades of sunflower oil, from
high-linoleic acid, to high-oleic acid (monounsaturated), to
intermediate types. Oleic acid, being monounsaturated, has
higher oxidative stability. In addition, the reduction in linoleic
acid means that the need for hydrogenation to reduce polyun-
saturated oils and the formation of trans-fatty acids is reduced.
Trans-fatty acids are nutritionally undesirable and similar to
saturated fats in their effects. Examples of plant species, which
have been bred with increased oleic acid, include sunflowers
(high oleic acid and NuSun), linseed (solin, Linola), cot-
tonseed, soybean, rapeseed (canola, Monola), safflower, and
mustard.
 THE OILSEEDS | Oilseeds: Overview 223

Table 2 The fatty acid components and quantities in the major oilseed crops (amounts in rounded percentages)

Fatty acid Name Palmitic Palmitoleic Stearic Oleic Linoleic Linolenic Arachidic Gadoleic Behenic Erucic

Symbol C16:0 C16:1 C18:0 C18:1 C18:2 C18:3 C20:0 C20:1 C22:0 C22:1

Oilseed Type

Soybean Normal 814 <0.2 26 1730 4859 411 <0.6 <0.5 <0.7 <0.3
Soybean High oleic acid 6 3 84 2 4
Cottonseed Normal 2126 <1.2 23 1522 4758 <0.5 0.20.5 <0.1 <0.6 <0.3
Peanuts Normal 8.3 3.1 56 26 2
Rapeseed Low erucic acid 27 <0.6 13 5170 1530 514 0.21.5 0.15 <0.6 <2.0
High erucic 1.56 <3 0.53 860 1123 513 <3 315 <2 260
acid
High oleic acid 34 23 6376 1325 23 12 <0.6 <0.2
Sunflower Linoleic acid 58 <0.3 27 1440 4874 <0.3 0.10.5 <0.3 0.31.5 <0.3
Mid-oleic acid 45 34 5075 2030 <1
Oleic acid 25 <0.1 37 7591 217 <0.3 0.20.5 0.10.5 0.51.5 <0.3
Sesame Normal 812 <0.2 56 3642 4248 0.30.4 0.30.6 <0.3 <0.3
seed
Linseed Normal 57 <0.3 34 1920 1417 5261 <0.5 <0.6
Low linolenic 6 34 15 73 23
acid
Safflower Linoleic acid 58 <0.2 23 822 6883 <0.1 0.20.4 <0.3 <1.0 <1.8
Oleic acid 36 <0.2 13 7084 920 <1 0.30.6 <0.5 <0.4 <0.3
Mustard B. carinata 410 <2 823 1522 1827 <2 2050
seed B. nigra 27 <2 1027 1522 1127 3345
B. juncea 310 13 1564 1428 924 13 <40

Source: Adapted from Codex Alimentarius Commission, Codex Stan 210-1999, Amend. 2013 and other literature values.

Other changes of significance to oilseed crops have been the Cottonseed

development of genetically modified plants with specific appli-
Cotton is basically grown for its fiber, but the seed has several
cations other than for edible oils. The most obvious example of
important components consisting of oil (16%), protein (37%),
these is soybean in which large numbers of products are in
hull (37%), and linters (10%). Also, within the kernel is a
development including those containing antibodies, altered
phenolic compound called gossypol,  1% of the seed weight:
amino acid profiles, and zero lipoxygenase. Rapeseed is also
toxic to humans and monogastric animals. Cottonseed oil,
in the process of undergoing numerous transformations to
prior to refining, is red due to the residual gossypol and asso-
produce products for polymers and detergents, inks, cosmetics,
ciated products. Increased seed oil content has been a second-
pharmaceuticals, lubricants, plasticizers, and resins.
ary objective to fiber production although the value of the oil is
significant. Cottonseed oil contains 4758% linoleic acid. To
make the oil more stable for cooking and to reduce the need for
hydrogenation, the Commonwealth Scientific and Industrial
The Crops
Research Organisation (CSIRO) in Australia have developed an
Soybean Seed alternative cottonseed oil. By switching off genes that convert
oleic acid to linoleic acid, they have been able to produce
The seeds are rich in protein, mainly globulins, which make up
high-oleic acid (73%), low-linoleic acid (5%) oil with
90% of the total proteins and 36% of the seed weight. Ten
increased oxidative stability. Additionally, they have found
percent of the seed weight is carbohydrate, mainly sugars, and
that it is possible to alter the proportions of saturated fatty
3% is starch. The seed is relatively low in oil compared with
acids, palmitic and stearic, which provide the solid properties
other oilseed crops at  20%. Following oil extraction, heat
necessary for making margarine.
treatment is used to inactivate enzymes in the high-protein
meal that otherwise reduce the digestibility of the stock feed.
Traditional soybean oil is polyunsaturated with 4859% lino-
Groundnut or Peanut
leic acid (C18:2) and 411% linolenic acid (C18:3) (Table 2).
Breeding has produced cultivars with various fatty acid profiles; Peanuts are a valuable source of oil and also produce a high-
in particular, high-oleic acid soybean oil, which contains up to protein meal for stock feed. Increased yields are an important
84% oleic, has less saturated (palmitic) acid and is more stable breeding priority, particularly in developing countries where
for cooking purposes. The meal may also be used for industrial yield is poor. Increased oil content is also desirable as peanuts
purposes such as cosmetics, paints, and adhesives and various have a wide range from 40% to >60% in wild types. Reduction
edible purposes. in linoleic acid produces oil with a better oxidative stability.
224 THE OILSEEDS | Oilseeds: Overview
Meal protein is high at 46%, but the amino acid profile is
lacking in sulfur-containing amino acids particularly methio-
nine. The components of whole peanuts are protein (26%), oil
(41%), and carbohydrates (24%). The main product is the oil
that has 80% unsaturated fatty acids making it a nutritionally
favorable oil. Additionally, the level of oleic acid and palmitic
acid makes it stable with good keeping qualities for cooking
and in food mixtures. Plant selection has seen the development
of new cultivars with high oleic acid levels of up to 60% and a
subsequent reduction in linoleic acid content. Fatty acid pro-
files are influenced significantly by environmental conditions
with high temperatures favoring high oleic acid contents.
Rapeseed
Traditional rapeseed oil characteristically contains high levels
of erucic acid, found to have detrimental effects on the myo-
cardial muscle of rats fed the oil. The value of the meal was also
low due to the presence of sulfur-containing compounds called
glucosinolates, found to have deleterious effects on the thyroid
gland of monogastric animals. Through plant breeding, major
changes have been achieved to traditional rapeseed by increas-
ing yield and oil while simultaneously increasing meal protein
content. The development of cultivars with very low levels of
glucosinolate in the meal has also been achieved. With the
dramatic change in rapeseed from traditional types to those
grown today, new terms have been used to discriminate
between the types. Canola is recognized under ISO 5725,
Codex Alimentarius, and ISTA as cultivars of rapeseed with
<2% erucic acid in the oil and < 30 mmol of aliphatic glucosi-
nolates in the meal. European nomenclature to describe
cultivars of rapeseed includes low-erucic acid rapeseed and
Colza, which is the French name for B. napus rapeseed in
general. The seed oil is the main value of the crop yielding
42% oil, while the meal contains 35% protein. Current canola
cultivars have very low erucic acid and saturated fatty acid
levels and a good proportion of oleic (C18:1), linoleic
(C18:2), and o-3 linolenic (C18:3) fatty acids. The balanced
fatty acid profile makes rapeseed ideal for mayonnaise and
salad dressings and a wide range of applications in the bakery
and confectionary industries. New high-oleic acid/low-
linolenic acid types have improved oxidative stability for cook-
ing and longer shelf life. Alternatively, high-erucic acid oil is
utilized for industrial purposes such as cutting oils. Erucic acid
is extracted from high-erucic acid rapeseed and converted to
erucamide for plastic manufacture.
Sunflower
Sunflower is grown for the seed oil, which is  80% of the seed
value. The dehulled meal has 2842% protein. The oil is highly
considered due to its low linolenic acid value and therefore
oxidative stability for cooking, salad oil, and margarines. Her-
itability of high oil is reliable, and it has been possible to
increase oil contents from 30% in early types to over 50% in
recent years. Sunflower cultivars with a range of fatty acid pro-
files have been developed including high oleic acid (7591%)
with reduced levels of linoleic acid (217%) and high levels of
alpha-tocopherol (vitamin E), providing the market with very
stable monounsaturated oil for cooking purposes. Mid-oleic
acid cultivars are also available, such as NuSun, with satu-
rated fatty acid levels of 8% and only 2030% linoleic acid. The
meal has high protein content and is used in animal feed for
livestock and poultry. A small percentage of the crop is used for
nonoilseed production for confectionary purposes.
Sesame Seed
Sesame seeds are used intact or as oil and meal. The seeds
generally have high oil content of 50% and  25% protein,
although the oil content can vary between 40% and 60%. The
fatty acid composition also shows a large range (Table 2).
A high oleic acid (40%) and linoleic acid (45%) content
makes the oil nutritionally beneficial. The unsaponifiable frac-
tion of the oil contains sesamin and sesamolin that during the
refining process form sesamol and sesaminol. These are strong
antioxidants that give the oil exceptional resistance to oxida-
tion and rancidity. The oil-free meal is high in protein
(3450%), depending on the variety, and has a favorable
amino acid profile with high methionine and low lysine con-
tent. Today, the seed is used for human consumption on bread
rolls, health food, and confectionary.
Linseed
Due to its high iodine value, linseed oil has been used primar-
ily for industrial purposes, such as linoleum floor covering,
with a high level of unsaturated fatty acids making the oil very
reactive and resulting in a short shelf life. Low linolenic acid
cultivars have introduced linseed to the edible food market. In
1994, the Flax Council of Canada developed the term solin to
describe linseed with < 5% linolenic acid. The original hybrid-
ization work was carried out by the CSIRO in Australia with the
release of two Linola cultivars in 1992 under the Plant Variety
Rights scheme. Linola 947 was the first solin cultivar registered
in Canada. Solin cv. Linola 989 has been reported as 46% oil
(dry basis) and 34% protein. Linseed meal has a high crude
protein value but low lysine levels. It also has a high level of
soluble fiber, called mucilage, that is indigestible to nonrumi-
nants and reduces the energy value of the meal. Linseed is
traded at 40% oil although the oil can range significantly
depending on growing conditions.
Safflower
Safflower was originally domesticated for its flowers: red
carthamin dye was extracted for coloring food and cloth.
Early cultivars were unsuitable for commercial development
due to low oil content of 30%. Breeding programs have since
increased oil contents and altered the fatty acid profiles to take
it from an industrial product to the edible oil market with
modern cultivars containing 34% or more oil. Traditional
high-linoleic acid types contain 6883% linoleic acid and
822% oleic acid (Table 2). Through plant breeding, high oleic
acid levels (7084%) have been achieved. Due to the wide
range of environmental conditions from countries as diverse
as Australia, India, and China, fatty acids may vary consider-
ably. The meal from safflower seed is also valued as a stock
feed. The protein content of the meal remains relatively low in
relation to other oilseeds at only 23%.
 THE OILSEEDS | Oilseeds: Overview 225

Cottonseed Peanuts Linseed

Mustard seed Rapeseed Safflower seed

Sesame seed Soybeans Sunflower seed

1000

100
Production (Mt)

10

0
05

06

07

08

09

10

11

12
20

20

20

20

20

20

20

20

Year
Figure 1 Global production of major oilseed crops (log10) from 2005 to 2012 in million tonnes (Mt). Adapted from Food and Agricultural Organization
of the United Nations Statistics.

Mustard Seed
Typically, mustard oil is high in erucic acid (2050%) and the
meal is high in glucosinolate compounds (50150 mM g 1), Table 3 World production of the major oilseed crops for 2011 and
2012
providing pungency to mustard for condiments but reducing
the palatability of the meal for stock feed. With plant breeding, Area harvested (Mha) Production (Mt)
very low levels of erucic acid have been achieved, and current
aims are to develop cultivars that are more closely related to 2011 2012 2011 2012
LEAR cultivars, with increased oleic acid content.
Soybeans 104 107 262 253
Rapeseed 34 34 63 65
Cottonseed NA NA 49 47
Production and Trade Groundnuts 25 25 40 41
Sunflower seed 26 25 41 38
For the purpose of this article, fruit oils will not be discussed Sesame seed 9 8 5 4
although production is significant. Palm oil fruit (from which Linseed 2 3 2 2
palmitic acid derives its name) production in 2012 exceeded Safflower seed 0.8 1.0 0.7 0.8
50 Mt, second only to soybean. Oilseed production from 2005 Mustard seed 0.7 0.6 0.6 0.5
to 2012 is shown in Figure 1 and the area grown and yields of NA, not available.
the various crops in 2011 and 2012 in Table 3. Source: Adapted from FAOSTAT | FAO Statistics Division 2014 | 03 February 2014.
226 THE OILSEEDS | Oilseeds: Overview

Soybean 2012. Other exporters included Ukraine, Hungary, and France.

Major importers include Turkey (0.9 Mt), the Netherlands
The global oilseed market is dominated by soybean with a total
(0.6 Mt), France (0.4 Mt), Germany (0.4 Mt), and Spain
of 242 Mt produced in 2012, making up almost 50% of the
(0.3 Mt).
total oilseed production (Table 3). The United States of
America, in 2012, produced 29% of world soybean produc-
tion, with Brazil (24%), Argentina (16%), India (10%), and
China (6%) also producing significant amounts. Not unex- Sesame
pectedly, the United States is the largest exporter followed by
World production of sesame seed has increased from 3.17 Mt
Brazil and Argentina.
in 2001 to over 4 Mt in 2012. All but a small part of total world
production is from developing countries including Myanmar
Rapeseed (0.6 Mt), India (0.6 Mt), China (0.6 Mt), and the United
Republic of Tanzania (0.5 Mt). India (0.4 Mt), Ethiopia
Rapeseed is now rated in second position in world production
(0.3 Mt), Nigeria (0.1 Mt), and Sudan (0.1 Mt) are the largest
of oilseeds with 65 Mt produced in 2012. There has been a
exporters of sesame. Major importers were China (0.4 Mt),
continual increase from 2005 to 2012 (Figure 1) in part due to
Japan (0.2 Mt), Turkey (0.1 Mt), Vietnam (0.1 Mt), and Repub-
increased production in Canada. Canada is now the largest
lic of Korea (0.1 Mt).
producer of rapeseed with 15.4 Mt per annum or 24% of the
world production. China is second with 14 Mt with India,
France, Germany, and Australia also being major producers.
Canada is also the major exporter of rapeseed (canola) with Linseed
exports of nearly 8 Mt in 2011, while France and Australia
World production of linseed has shown a gradual decline since
exported around 1.6 Mt each. Germany is the biggest importer
the 1980s despite a peak in 1999 of almost 3 Mt. The produc-
of rapeseed (canola) importing 3.2 Mt in 2011, with Japan
tion has continued to fall between 2005 (2.8 Mt) and 2012
(2.3), the Netherlands (2.3), Belgium (2.2), Mexico (1.6),
(2.1 Mt) (Figure 1). The total production of 1.7 Mt in 2007
and China (1.3) also being major importers.
was the lowest level of production in the previous 46 years.
Canada, the largest producer, decreased production from
Cottonseed 1.0 Mt in 2005 to 0.5 Mt in 2012. The major producers of
linseed are Canada, Russia, China, Kazakhstan, the United
The major purpose of cotton is for textile use. Production of
States, and India. Canada is the major exporter with
cottonseed for oil has continued to increase in the last 20 years
359 000 T exported in 2011. Russia, Belgium, and Kazakhstan
(Figure 1) although it has recently slipped back from the
are also major exporters. Belgium, the United States, Germany,
second largest oilseed crop to third place after rapeseed. Global
and China are major importers.
production is 47 Mt with China being the largest producer at
14 Mt per annum. Other major producers include India
(10.8 Mt), United States (5.1 Mt), Pakistan (3.2 Mt), Brazil
(3.3 Mt), Uzbekistan (2.0 Mt), Australia (1.7 Mt), and Turkey Safflower
(1.3 Mt). Australia is the leading exporter of cottonseed while
Production of safflower has also been relatively stable over
Mexico, Spain, Japan, Italy, Korea, and the United States are
many years, falling from 900 kt in 1985 to 600 kt in 2002 but
large importers.
rising again in 2012 back to 800 kt. This failure to increase may
reflect the increased demand for monounsaturated oil in place
Groundnuts or Peanuts of polyunsaturated oils. The major safflower-producing coun-
tries in 2012 are Mexico (258 kt), India (152 kt), Kazakhstan
A global groundnut production of 41 Mt is led by China (127 kt), Argentina (108 kt), and the United States (81 kt). The
(17 Mt or 41%) with India (14%), Nigeria (8%), the United United States and Australia are major exporters of safflower, but
States (7%), and Myanmar (3%) also being significant pro- the United States is also a net importer of safflower oil. Japan is a
ducers. The major exporters of groundnuts include India major importer.
(0.8 Mt), Argentina (0.2 Mt), and the United States of America
(0.1 Mt). Importers are the Netherlands (0.3 Mt), Indonesia
(0.1 Mt), Mexico (0.1 Mt), and Russian Federation (0.1 Mt).
Mustard
There has been no increase in world production of mustard in
Sunflower
recent years (Figure 1) from 565 kt in 2005 to 494 kt in 2012.
Sunflower production has continued to increase in recent Major producers in 2012 include Nepal (145 kt), Canada
years, in the period between 2005 and 2012, from 31 to (118 kt), Myanmar (68 kt), and Russia (41 kt). In 2011,
38 Mt (Figure 1). Ukraine is now the biggest producer with Canada exported 130 kt of mustard, while Ukraine (39 kt),
8.4 Mt in 2012, which is 22% of world production. The Russia (34 kt), Germany (21 kt), and Czech Republic (12 kt)
Russian Federation also produced 8 Mt in 2012. Argentina, were also significant exporters. The United States (68 kt), Ger-
China, and France are other major producers. Romania many (50 kt), Nepal (32 kt), France (25 kt), and Bangladesh
(1.2 Mt) and Bulgaria (1.0 Mt) were the major exporters in were the major importers in 2011.

Future Developments
Oilseeds play an increasingly important role in society, both as
an edible food product and for industrial uses. The benefits
include the valuable oil component and the secondary but also
useful meal that provides high-energy and nutritionally impor-
tant food. Breeders have been able to optimize the characteris-
tics of many of these crops to achieve higher levels of
production and to alter the products to suit specific require-
ments. These changes include improvements in nutritional
value, particularly with changes in fatty acid profiles and the
reduction in antinutritional components. Undoubtedly the
biggest changes will come in the future from genetic modifica-
tion in which long-term breeding programs can be dramati-
cally reduced to relatively short periods of time. Traditional,
breeding techniques rely on natural variability in plant popu-
lations to achieve change. Genetic engineering can create new
products previously out of the reach of established breeding
techniques. Engineering plants for specific aims or more suit-
able to environmental conditions can increase production.
Nutritionally, fatty acid profiles can be dramatically altered to
reduce saturated and trans-fatty acids and simultaneously
improve oxidative stability. Vegetable oils are already being
substituted, in many instances, for less environmentally
friendly petroleum products. The major competition to oilseed
crops in the future will only be from the ever-growing range of
additional oilseed types.
Exercises for Revision
Discuss the major oilseed crops of the world and their uses.
Which are the major producing and importing countries for
each species of oilseed?
Consider and discuss what changes have been made to
make oilseed crops more useful?
Evaluate the importance of fruit oils and how do they rate
against oilseed crops?
Compare oilseeds that are used for oil with the ones that are
grown for their high-protein meal and consider why?
Exercises for Readers to Explore the Topic Further
Consider the changes in oilseed production over years and
the ability of breeders and producers to keep pace with the
demand of an increasing population.
THE OILSEEDS | Oilseeds: Overview 227
Are the same countries always the largest producers or is
this changing? Why are some countries reducing
production, while others are increasing?
How are the uses of oil, meal, and oilseeds changing? Look
at the new applications for these products, which differ
from the original use for food.
What will be the impact of industrial uses, particularly
biodiesel to the food supply in the future?
See also: Agronomy of Grain Growing: Canola: Agronomy;
Soybean: Agronomy; The Legumes and Pseudocereals: Peanuts:
Overview; The Oilseeds: Cottonseed: Overview; Sunflower: Overview.
Further Reading
Appelqvist L-A and Ohlson R (eds.) (1972) Rapeseed: Cultivation, Composition
Processing and Utilization Amsterdam: Elsevier.
Benedict JH, Treacy MF, and Kinard DH (eds.) (1994) Vegetable Oils and Agrichemicals
Memphis, TN: Cotton Foundation and National Cottonseed Products Association.
Bennett M (1998) Sesame seed. In: A Handbook for Farmers and Investors Australia:
Rural Industries Research and Development Corporation.
FAO (2003) FAO Statistical Database. Rome: Food and Agriculture Organization of the
United Nations.
Hui YH (ed.) (1996) Baileys Industrial Oil and Fat Products, 5th edn., Series 3. New
York: Wiley ISBN: 047159427X.
Karleskind A (1996) Oil and Fats Manual. France: Lavoisier Publishing.
KeShun L (1997) Soybeans: Chemistry, Technology and Utilization. New York:
Chapman and Hall, ISBN: 0-8342-1299-4.
Kimber DS and McGregor DI (eds.) (1995) Brassica Oilseeds, Production and
Utilisation Wallingford: CAB International.
Robbelen G, Downey RK, and Ashri A (1989) Oil Crops of the World. New York:
McGraw-Hill.
Weiss E (2000) Oilseed Crops, 2nd edn. Malden, MA: Iowa State Press.
Woodroof JG (1983) Peanut Production, Processing, Products, 3rd edn.
Westport, CT: AVI.
Relevant Websites
http://www.amsoy.org American Soybean Association.
http://www.aocs.org American Oil Chemists Society.
http://www.apps.fao.org Food and Agriculture Organization.
http://www.australianoilseeds.com Australian Oilseed Federation.
http://www.australianoilseeds.com 7th International Safflower Conference.
http://www.canola-council.org Canola Council of Canada.
http://www.codexalimentarius.net Codex Alimentarius Commission.
http://www.cottonseed.com National Cottonseed Products Association.
http://www.flaxcouncil.ca Flax Council of Canada.
http://www.grainscanada.gc.ca Canadian Grain Commission.
www.sunflowernsa.com National Sunflower Association.
www.regional.org.au 10th International Rapeseed Congress.
http://www.rirdc.gov.au Australian Rural Industries and Research Foundation.
 Soybean: Overview
K Liu, Agricultural Research Service, US Department of Agriculture, Aberdeen, ID, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Soybean is an ancient Chinese grain and now a top oilseed
crop, with world annual production reaching more than
300 million tonnes.
Soybean is also a legume, having the ability to fix nitrogen
from air through its roots.
Soybean has the highest protein content (40%) and the
second highest oil content (20%) among food legumes. It
also contains an array of phytochemicals.
Soybean has a versatile use as food, feed, and industrial
material.
Soyfoods come in various types: traditional, modern
(Westernized), fermented, and nonfermented.
Soyfoods are not only nutritious but also healthy since
regular consumption of soyfoods can help prevent certain
chronic diseases.
The majority of soybean produced annually is currently
crushed into oil for edible consumption and defatted
meal mostly for animal feed. Only a small portion of
annual production is used for whole soy food or soybean
protein ingredients.
Learning Objective
To have a basic understanding of soybeans with respect to
its history, current production, agronomic traits, chemistry,
nutrition, processing, and various end uses
Introduction
From one of the five sacred grains in ancient China to one of
top cash crops, the soybean (Figure 1) has played a very
important role in nourishing humans and animals. For the
past several decades, progress has been made on many fronts
of soybean research, from production management to variety
development, from genome mapping to compositional modi-
fication, from processing to new product development, from
health benefits to identification of key responsible compo-
nents, and from animal nutrition to new industrial uses.
Since there are so many topics to be covered on soybeans, it
is impossible to give a detailed account in this article. Readers
are advised to refer to several specific, topical chapters on
soybeans as well as further reading under each article.
Production
For the past six decades, the soybean has emerged as one of the
most important agricultural crops in the world, with a steady
increase in annual production. In 1955, the world soybean
production was only about 20 million tonnes (MT). In 2014,
global production increased to about 311 MMT (Figure 2).
228 Encyclopedia of Food Grains, Second Edition
During the same period, soybean production in the United
States increased from about 9 to 106 MMT and in Brazil and
Argentina from virtually negligible amounts to 94 and
55 MMT, respectively, while soybean production in China
went up and down within a small range of 1015 MMT.
These differential changes have made the United States, Brazil,
and Argentina the top three soybean producers and China
from the worlds number 1 prior to the 1950s to the worlds
number 4 soybean producer at the present time.
There are several reasons for the rapid and sustainable
worldwide growth in soybean production. First, soybeans
have many possible end uses, including as human food, ani-
mal feed, and industrial materials. Second, advances in plant
breeding and production management have led to a steady
increase in production yields in the field and improvements
in the seed quality. This is particularly true in recent years when
application of crop biotechnology has resulted in the develop-
ment of new soybean varieties that are resistant to certain
diseases and pests, tolerant to herbicides, and/or improved in
seed composition. Third, the soybean is capable of producing a
higher amount of edible protein than any other crops per
hectare of land while producing a significant amount of edible
oil. Fourth, like other members of the legume family, soybeans
have an ability to fix nitrogen due to its characteristic symbiosis
with root bacteroids. This feature makes the soybean a good
rotational crop. Fifth, recent medical discoveries have linked
the consumption of soyfoods to the prevention and treatment
of chronic diseases such as heart diseases, cancer, and bone
diseases. This new development helps promote soybean utili-
zation as food. Finally, growth and globalization of world
economy and trades have been another important driving
force. We are seeing a rapid interchanging trend with respect
to soybean utilization between China and the rest of the world.
In China, where soybeans originated and traditional soyfoods
are popular, rapid economic development in recent years has
led to increased demand for soybeans as refined oil and animal
feed. In the rest of world, particularly in North America and
Europe, where historically soybeans are crushed for edible oil
and animal feed, interest in soyfoods and soy-enriched food
has experienced a phenomenal rise over the past 2 decades.
Origins
Historical and geographic evidence indicates that soybeans
originated in northern China and that its cultivation in the
region started approximately 5000 years ago. The soybean was
considered one of the five sacred grains, along with rice, wheat,
barley, and millets. It was repeatedly mentioned in later
records. The Chinese gradually developed many nutritious
foods out of soybeans, including soy milk, tofu, soy sprouts,
fermented soy paste, and soy sauce (Figure 3). Soybean culti-
vation and soyfood preparation methods were gradually intro-
duced into Japan, Korea, and other neighboring countries
about 1100 years ago. People in these countries not only
http://dx.doi.org/10.1016/B978-0-12-394437-5.00028-0
 THE OILSEEDS | Soybean: Overview 229

accepted the Chinese way of preparing soyfoods but also cre-
ated new soyfoods. Japanese natto and Indonesian tempeh are
two good examples. All these foods are now known as tradi-
tional soyfoods.
The soybean and soybean products were introduced into
Europe during the Age of Discovery (sixteenth to seventeenth
centuries) and into the North America during the mid-
eighteenth century. Yet, large-scale official introduction into
the United States did not occur until late nineteenth century
and early twentieth century, during which the US Department
of Agriculture developed a national soybean germplasm col-
lection, and thousands of new varieties were collected, mostly
from China. Currently, the collection contains over 20 000
strains, serving as a primary genetic source for soybean research
and variety development.
Agronomic Characteristics
The soybean belongs to the family Leguminosae. The culti-
vated form, Glycine max (L.) Merrill, grows annually. Its plant
is bushy, with a height between 0.2 and 2.0 m, and produces a
fruit having three to five hairy pods in a cluster (Figure 4). Each
pod, 38 cm long, contains two to four seeds that are spherical-
to oval-shaped. Most seeds are yellow, but they can also be
green, dark brown, purplish black, or black.
Most mature plant seeds are made of three basic parts: the
seed coat, embryo, and one or more food storage structures
(such as the endosperm). The soybean seed, however, like
other legume seeds, is essentially devoid of endosperm and
consists of only a seed coat and a large, well-developed

Figure 3 Traditional soyfoods: soy milk, soy sauce, soy milk film, soy
Figure 1 Soybean seeds. sprouts, tofu, and okara (soy fiber).

300 Total World

USA
250
Brazil

Argentina
200
Million tonnes

China

150 India

Paraguay
100 Canada

Others
50

0
1955 1960 1970 1980 1990 2000 2010 2014
Production Year
Figure 2 Soybean production around the world. Data extracted from oil crops outlook, Economic Research Service, US Department of Agriculture,
Washington, DC.
230 THE OILSEEDS | Soybean: Overview
Figure 4 Harvesting soybeans in the field. Courtesy of the United
Soybean Board.
embryo. The seed coat is marked with a hilum (also known as
seed scar) that varies in color. The embryo contains two equal
cotyledons and three other structural parts: radical, hypocotyl,
and epicotyls. The radicle and hypocotyl, together known as
embryonic axis or germ, are located under the seed coat at one
end of the hilum, just below the micropyle, which is a tiny hole
formed by the integuments during seed development. The
epicotyl is tucked between the pair of cotyledons. During
germination, the radicle becomes the primary root, whereas
the hypocotyl lifts the cotyledons above the soil surface.
The epicotyl is the main stem and growing point. Cotyledons
function as a nutrient reservoir. In general, cultivated soybeans
constitute approximately 8% hull, 90% cotyledons, and
2% germ.
The soybean plant has two major growth stages: the vege-
tative stage and the reproductive stage. The period between the
emergence and the appearance of the first flower is the vegeta-
tive stage. This usually takes 68 weeks. The soybean plant is
photoperiod-sensitive and makes the transition from vegeta-
tive to flowering stages in direct response to day length. Most
soybean varieties begin flowering soon after the day length
begins to shorten. Because of this feature, latitude plays a
major role in the adoption of soybeans to various geographic
regions. Following the period of vegetative growth, the soy-
bean plant enters the reproductive stage in which maxillary
buds develop into flower clusters of 235 flowers each. This
is followed by pod development, seed development, and mat-
uration. The whole stage lasts from 7 to 12 weeks.
Like other farm crops, soybean plants are vulnerable to a
wide range of bacterial, fungal, and viral diseases and parasites
(such as cyst nematodes). They are also exposed to weeds and
other environmental stresses, such as saline or alkaline soils,
drought, and chilling temperatures. Rotating crops, spraying
chemicals, and choosing resistant cultivars have been the pri-
mary tools used by farmers to control diseases, pests, and
weeds and manage other stresses. However, the wide use of
pesticides, herbicides, and other chemicals has caused environ-
mental concerns. Fortunately, advances in plant breeding and
genetic engineering have made the biological control quite
possible. New soybean varieties with herbicide tolerance,
insect resistance, and disease resistance have been developed
and successfully commercialized. Among them, glyphosate-
Figure 5 General composition of soybean seeds. Courtesy of the United
Soybean Board.
tolerant soybeans and Bt soybeans, both transgenic in nature
(also known as genetically modified organism), have been
widely adapted in the field.
Chemistry and Nutrition
General Composition
Soybeans are known as yellow jewel or miracle beans,
because of their unique chemical composition (Figure 5). By
average, on a dry matter basis, mature soybeans contain about
20% oil, 40% protein, and 5% ash and the remaining 35% is
total carbohydrate. With this composition, the soybean has the
highest protein content among cereal and other legume spe-
cies. Protein content in other legumes ranges between 20% and
30%, whereas those in cereals range between 8% and 15%. The
soybean also has the second highest oil content among all food
legumes (after the peanut). Because of this, the soybean is
considered an oilseed as well. Other minor components
found in soybeans include minerals, vitamins, and various
biologically active substances.
The majority of soybean protein is storage protein. Based
on solubility patterns, soybean seed proteins are divided into
albumins and globulins. Albumins are soluble in water,
whereas globulins are soluble in a salt solution. Soybean glob-
ulins are the primary portion of total seed proteins and can be
further divided into glycinin and conglycinin. Due to differ-
ences in composition and structure, the two exhibit differences
in nutritional quality (amino acid composition) and func-
tional properties (such as gelation and thermal stability).
During seed development, soybeans store their lipids in an
organelle known as oil bodies, mainly in the form of triglycer-
ides. During processing, components extracted from soybeans
by hexane are classified as crude oil. Triglycerides are the major
component of crude oil, while minor components include
phospholipids, unsaponifiable material, free fatty acids, and
trace metals. Soybean oil typically contains about 11% palmi-
tic, 4% stearic, 23% oleic, 53% linoleic, and 8% linolenic acids.
With this fatty acid composition, soybean oil remains liquid
over a wide range of temperatures.
Soybean carbohydrates can be grouped into insoluble and
soluble ones. The insoluble carbohydrates include cellulose,
hemicellulose, pectin, and a trace amount of starch. They are

structural components found mainly in cell walls. Soluble
carbohydrates include measurable amounts of di- and oligo-
saccharides, with sucrose in the range of 2.58.2%; raffinose,
0.10.9%; and stachyose, 1.44.1%, and trace amounts of
monosaccharides, such as glucose and arabinose. The oligosac-
charides are nonreducing sugars, containing fructose, glucose,
and galactose as three or four units, linked by beta-fructosidic
and alpha-galactosidic linkages.
The content of major minerals in soybeans ranges from
0.2% to 2.1%. Among them, potassium is in the highest con-
centration, followed by phosphorus, magnesium, sulfur, cal-
cium, chloride, and sodium. The minor minerals present in
soybeans are silicon, iron, zinc, manganese, copper, molybde-
num, fluorine, chromium, selenium, cobalt, cadmium, lead,
arsenic, mercury, and iodine. Their content ranges from 0.01
to 140 ppm. Soybeans also contain both water-soluble and
oil-soluble vitamins. The water-soluble vitamins present in
soybeans include thiamin, riboflavin, niacin, pantothenic
acid, and folic acid. The oil-soluble vitamins present in soy-
beans are vitamins A and E with essentially no measurable
vitamins D and K.
Biological Active Components
There are an unusually large number of heat-stable and heat-
labile substances that are naturally found in soybeans. These
compounds, although present in a trace amount, are capable of
eliciting diverse nutritional, biological, and physiological
responses in humans and animals. These biological and phys-
iological responses can be adverse or beneficial to human and
animals, and as such, some substances are known as anti-
nutritional factors, while others are referred to as phytochem-
icals or nutraceuticals. Sometimes, the same substances may
exert both adverse and beneficial effects, depending on the
dosage and specific application. Due to page limitations, only
a few key compounds are briefly described in the succeeding
text.
Proteinase inhibitors present in soybeans fall into two
types: Kunitz trypsin inhibitor and BowmanBirk inhibitor.
The significance of soybean trypsin inhibitors lies in their
nutritional implications toward both humans and animals.
Unless inactivated by heat, these inhibitors can cause growth
depression by reducing the digestibility of proteins.
Lectin, also known as hemagglutinin, is another antinutri-
tional factor that is of protein nature found in soybeans. It
possesses a remarkable ability to agglutinate erythrocytes and
other types of cells. As demonstrated by animal studies, soy-
bean lectin is linked to an enlargement of the pancreas, a
lowering of blood insulin levels, an inhibition of disacchari-
dase and proteases in the intestines, and degenerative changes
in the liver and kidneys, in addition to growth inhibition.
Phytate is the calciummagnesiumpotassium salt of ino-
sitol hexaphosphoric acid, which is commonly known as phy-
tic acid or phytin in the literature. As in most seeds, phytate is
the principal source of phosphorus in soybeans. Its content
ranges from 1.00% to 1.47% on a dry matter basis. This value
represents a little more than 50% of the total phosphorus in
soybean seeds and 6070% of total phosphorus in defatted
soybean meal. Phytate P is nutritionally unavailable to mono-
gastric animals. Supplementation with inorganic P into
THE OILSEEDS | Soybean: Overview 231
soy-based feed has been a practical way to address the problem.
Excess P due to unavailability or oversupplementation in ani-
mal feed is excreted in feces, raising environmental concerns.
Among soybean carbohydrates, raffinose and stachyose gar-
ner the most attention, mainly because human and monogas-
tric animals cannot digest these sugars well due to the lack of a
specific, endogenous alpha-galactosidase. As a result, their
presence has been linked to flatulence and abdominal discom-
fort associated with soybean consumption.
Phospholipids are polar lipids, constituting 0.51.5% of
the soybean seed and 13% of crude soybean oil. The com-
mercial product of phospholipids is known as lecithin, a major
by-product of the soybean oil refining process. Phospholipids
are good emulsifying agents, soluble in alcohol and insoluble
in acetone. In living tissues, they are the major components of
cell membranes.
Isoflavones belong to a group of compounds that share a
basic structure consisting of two benzyl rings joined by a three-
carbon bridge that may or may not be closed in a pyran ring.
The soybean is unique in that it contains the highest amount of
isoflavones, being up to 3 mg g 1 dry weight. Among all the
health-promoting components, isoflavones are thought to be
most responsible for many of the hypothesized health benefits
of soyfoods and thus have gained the most attention in the
scientific community.
Soybeans contain 0.10.3% saponins, which are composed
of sugars bound to alkaloid, steroid, or triterpene compounds
and have detergent surfactant properties. Saponins have blood
cholesterol-lowering properties rendered by their binding of
cholesterol. The bound cholesterol is then passed to the colon
and excreted.
Phytosterols are lipid-like compounds found in plants.
Soybeans, rapeseeds, and coniferous trees are the three major
commercial sources of phytosterols. The total phytosterol con-
tent of soybeans is estimated at 0.30.6 mg g 1. Soybean ste-
rols are obtained during oil processing as by-products of
vitamin E manufacturing. Although phytosterols are structur-
ally related to cholesterol, they have been clinically proven to
reduce blood cholesterol in our bodies.
Nutritional Values
In addition to their high content in soybeans, soybean oil and
protein are both of high quality. Like other edible oils, aside
from culinary roles, soybean oil provides us with calories,
essential fatty acids, and fat-soluble vitamins. Since soybean
oil contains very high percentages of unsaturated fatty acids, it
is an excellent source of essential fatty acids.
Soybean protein contains all the identified essential amino
acids for animals: isoleucine, leucine, lysine, methionine,
cysteine, phenylalanine and tyrosine, threonine, tryptophan,
valine, and histidine. Most of these amino acids are present in
amounts that closely match those required for humans and
other animals. The only potentially limiting amino acids are
sulfur-containing ones, primarily methionine and cysteine.
Yet, such deficiency is increasingly considered less problematic,
with respect to human nutrition. Protein quality depends on
the amino acid composition of the protein, amino acid
requirements for humans or animals fed with the protein, the
active amounts of antinutritional factors, digestibility of the
232 THE OILSEEDS | Soybean: Overview
protein, and the method for assaying protein quality. In the
earlier days, protein quality was assayed by the protein effi-
ciency ratio, which was measured by feeding the protein of
interest to rats. Since rats have a much higher need for sulfur-
containing amino acids than humans, soybean protein quality
was underestimated. Today, the protein digestibility-corrected
amino acid score (PDCAAS), a newer method, has been
adopted. Since it compares the pattern of essential amino
acids in a protein with the requirements that humans have
for essential amino acids and then adjusts for how well that
protein is digested, PDCAAS measures how limiting each indi-
vidual amino acid is in a protein. Under this new assay
method, soybean protein has a score very close to 1, the highest
rating possible, which is the same rating for animal proteins,
such as egg white and casein.
The deficiency in sulfur-containing amino acids associated
with soybean proteins can readily be corrected by a simple
change in the dietary patterns in humans. For example, since
cereal grains are normally rich in sulfur-containing amino
acids (but poor in lysine), one easy solution has been to
combine soyfoods with cereals. This dietary pattern has in
fact been practiced in the Orient throughout the history of
soyfood consumption even without much understanding of
food chemistry and human nutrition. For monogastric ani-
mals, suboptimal concentrations of cystine and methionine
in soy-based feed are effectively addressed by methionine
supplementation.
Health Benefits
Historically, the soybean has been primarily identified with its
high protein quantity and quality. Yet, for the past several
decades, medical research has linked consumption of soy to
the prevention and treatment of several chronic diseases and
implicated soybean proteins, isoflavones, and other phytochem-
icals as contributing factors. For example, a mega study con-
ducted in 1995 showed that soybean protein can lower total
cholesterol (9.3%), low-density lipoprotein (LDL) cholesterol
(12.9%), and triacylglycerols (10.5%), with a small increase in
high-density lipoprotein (HDL) cholesterol. Formal recognition
of the cholesterol-lowering properties of soybean protein came
in 1999 when the US Food and Drug Administration approved a
health claim for the cholesterol-lowering effects of soybean
protein. This ruling has set off a rush by many US mainstream
food companies to enter the soyfood market, helped increase
the awareness of soyfood products, and created an incentive for
food processors to incorporate soybean protein ingredients into
many types of existing food. As a result, the soyfood industry in
the United States and around the world has experienced phe-
nomenal growth and opportunity.
Soybean Processing
Simply boiling whole soybeans for direct use is mostly limited
to immature soybeans. For any effective use, soybeans have to
be processed into food, dietary ingredients, or coproducts.
Narrowly defined, soybean processing includes traditional pro-
cessing, modern processing, and alternative processing, all
Figure 6 Soybean oil and defatted meal. Courtesy of the United
Soybean Board.
starting with whole beans. Broadly defined, soybean proces-
sing also includes secondary processing, which starts with soy
ingredients or intermediate products, such as crude oil refin-
ing, soybean oil processing, soybean protein processing, and
coproduct recovering.
Modern Processing
Modern processing is basically a solvent extraction to produce
crude oil and defatted meal (Figure 6). The process consists of
preparation, extraction, and desolventization. Soybean prepa-
ration includes cleaning, drying, cracking, conditioning, and
flaking, with dehulling (before conditioning) as an optional
step. During solvent extraction, oil is extracted from the soy
flakes by an organic solvent (most often hexane). Two inter-
mediate products come out of the extractors: an oilsolvent
mixture and defatted soy flakes. Both need to be desolventized
to remove solvent. The oilsolvent mixture has about 2540%
hexane, which can be removed and recycled by an evaporator.
Defatted flakes contain about 30% hexane, and the choice of
desolventing equipment depends on the end use of desolven-
tized flakes (also known as defatted meal). For the majority of
meal used as animal feed, flakes are processed through a
desolventizertoaster, which uses hot steam to remove solvent
and inactivate heat-labile antinutritional factors, such as tryp-
sin inhibitors and lectins. This is to ensure optimal nutritional
value of meals. For food aimed at human consumption, it is
necessary to minimize heat denaturation during desolventiza-
tion of flakes to main high water solubility of soybean pro-
teins. This is carried out in a vapor desolventizing system or a
flash desolventizing system.
Crude oil contains variable amounts of nontriglyceride
compounds. In order to remove these impurities to produce
high-quality edible oil, crude oil needs to be subjected to a
series of refining operations. This includes degumming, neu-
tralization, bleaching, and deodorization. Degumming
removes phospholipids (also known as gums). The recovered
gums are the source of commercial lecithin. Neutralization
removes free fatty acids. Bleaching removes most colorants
and some residues from previous refining steps. Deodorization
is a steam stripping process to vaporize undesirable volatile
 THE OILSEEDS | Soybean: Overview 233

components. It is the final step after general processing. The Traditional Soyfoods
final product is refined, bleached, and deodorized (RBD) oil,
Over the course of several thousand years, long before the
commonly known as RBD oil. Yet, due to its high concentra-
modern processing emerged in the twentieth century, soybeans
tions of polyunsaturated fatty acids, RBD oil is prone to oxida-
had been processed into various soyfoods, known as tradi-
tion and off-flavor formation. It also lacks certain physical and
tional soyfoods. Since these foods were created in China and
functional properties. To overcome these limitations, addi-
neighboring countries, they are also known as Oriental soy-
tional processes (collectively known as soybean oil modifica-
foods. In general, traditional soyfoods are classified as non-
tion), such as hydrogenation, interesterification, winterization,
fermented and fermented. Nonfermented soyfoods include soy
and/or fractionation, may also be used.
milk, tofu, soy sprouts, yuba (soy milk film), okara (soy pulp),
immature soybeans, soynuts, and toasted soy flour (Figure 3).
Traditionally, soy milk is made in China by soaking, rins-
Alternative Processes ing, and grinding soybeans into slurry. This is followed by
filtering the slurry to separate the residue (okara) and cooking
There has been increased concern over the past several decades
the extract to become edible. As shown in Figure 7, the pro-
with the safety and environmental impact of hexane solvent-
cesses for making tofu, soy milk film, and okara are directly
based extraction. Several alternative oilseed milling processes
linked to soy milk production; tofu and soy milk film all start
have been available or under development. Screw pressing or
with soy milk making while okara is a by-product of soy milk
expelling is an old way of processing soybeans and other
making. To make tofu, soy milk is boiled for about 10 min,
oilseeds, which existed long before solvent extraction was
and a coagulant suspension is added to form curds, which is
developed. The method, also known as mechanical extraction,
then shaped and pressed into tofu.
has relatively low oil removal efficiency (leaving 48% residual
There are four major fermented soyfoods fermented soy
oil in the meal) and low production efficiency. The advantages
paste (miso), soy sauce, tempeh, and natto and two minor
are low initial cost and no solvent requirements.
fermented soyfoods: sufu (fermented tofu) and soy nuggets
Dry extrusion is another alternative processing method for
(fermented whole soybeans). They vary greatly in the methods
soybeans. It was developed in the 1960s to enable Midwestern
of preparation, microorganisms involved, and utilization.
US soybean growers to cook soybeans for use as livestock feed
While it takes less than a week to prepare tempeh and natto,
right on the farm where soybeans were produced. The process
the preparation of the remaining types of fermented soyfoods
generates friction as the sole source of heat to deactivate anti-
generally requires several months to complete. Except for
nutrients in soybeans. It does not remove any oil, though. Yet,
natto, which results from bacterial (Bacillus natto) fermenta-
when dry extrusion is combined with expelling, the through-
tion, all others are fermented mainly by fungi: soy paste, soy
put of the mechanical extraction is greatly improved. The pro-
sauce, and soy nuggets are all fermented by Aspergillus oryzae;
cess, known as extrudingexpelling, was developed in the
tempeh by Rhizopus oligosporus; and sufu by Mucor sp., Actino-
1980s and has become the primary choice for alternative pro-
mucor sp., and others.
cessing of soybeans.
Almost all nonfermented soyfoods are consumed primarily
Another alternative method for processing soybeans is
for nourishment. In contrast, the majority of fermented
aqueous extraction process. The method differs radically from
standard oil milling processes in that an aqueous system is
used to extract oil. Although the concept and general features
of the aqueous method appear attractive compared to the Whole soybeans
conventional hexane-based method, the comparatively low
oil yield and relatively high content of oil in the remaining Soaking
fractions have discouraged its commercial application.
Water Rinsing

Grinding

Soybean Utilization Filtrating Okara

The soybean is the most versatile grain in terms of its end use. Cooking
Broadly speaking, it can be used as human food, animal feed,
and industrial materials. At the present, the majority of the Soymilk
annual soybean production is crushed, and its oil is solvent-
extracted with hexane for edible use. The defatted soybean Coagulant Coagulating
meal is generally toasted and serves as feed for various farm
animals and fish. A very small proportion of annual soybean Pressing (optional) Whey
production is processed into many types of traditional and
modern soyfoods. In addition, both the oil and protein meal Tofu
have been explored for a broad range of industrial uses, includ- Figure 7 Outline of soy milk and tofu making. Adapted from Liu K
ing paints, plastics, and adhesives. Most notably, soybean oil (1997) Soybeans: Chemistry, Technology, and Utilization. New York:
has been used as a feedstock for producing biodiesel to replace Chapman & Hall, Figure 4.2, with kind permission of Springer Science &
petroleum-based transportation fuels. Business Media.
234 THE OILSEEDS | Soybean: Overview
soyfoods contain salt and/or by-products (such as lactic acid)
from a desirable fermentation. Both inhibit, delay, or slow
down spoilage of these products and allow them to have
a relatively long shelf lives. They are generally served as sea-
sonings or condiments in cooking or making soups and con-
tribute more in flavor than in nutrition to the diet. Tempeh and
natto are two exceptions, which have short shelf lives, are often
consumed as a part of the main meal, and contribute protein
and oil to the diet in addition to their characteristic flavor.
In commercial processing, most traditional ways of making
soyfoods are now modernized. For example, in making mod-
ern soy milk, several key steps are modified or added to reduce
beany flavor, increase production yield, and improve overall
product quality and consumer acceptance, even though the
basic principle is very similar to the traditional Chinese
method. The new steps include dehulling, blanching, fine
milling, centrifuging, formulation, fortification, homogeniza-
tion, thermal processing, and aseptic packaging.
Soybean Oil
Nearly all soybeans are processed for their oil and defatted
meal. With rare exceptions, crude soybean oil will have been
refined, bleached, and deodorized to produce a high-quality
RBD oil. About 85% of soybean oil is used for human con-
sumption. The rest is sold for biodiesel production or indus-
trial uses.
Soybean oil is the leading edible oil and widely used in
various food products. Among them are salad and cooking oil,
shortening, margarine, mayonnaise, and salad dressing. Other
applications include uses in canned foods, confectionery coat-
ings, pudding mixes, pancake and waffle mixes, macaroni and
cheese mixes, spaghetti sauces, dry breakfast cereals, frozen-
fried seafood, meat patties, pizza mixes, and others.
Since soybean oil contains relatively high proportions of
unsaturated linoleic and linolenic acids, it is susceptible to off-
flavor development. It also lacks some physical properties for
certain applications, such as shortening and margarine. To
increase oxidative stability and the melting point of soybean
oil, hydrogenation was commonly used until a few years ago.
The process chemically adds two hydrogen atoms to a double
bond of an unsaturated fatty acid in the oil. As a result, lower
unsaturated fatty acids are converted to saturated acids and
higher unsaturated fatty acid to lower unsaturated acids in
the oil. However, hydrogenation causes the formation of
trans-fatty acids. In recent years, the adverse health effects of
trans fat has been well established, causing US regulatory agen-
cies to call for reducing or even totally eliminating trans fat
from foods commercially prepared for human consumption.
This has created a great challenge for the food industry to find
alternatives for meeting different application needs.
One effective way to solve the trans fat issue, while keeping
the stability and functional properties of soybean oil required
for many edible applications, is through plant breeding and
genetic engineering. Already, several genotypes possessing
large variation in fatty acid composition have been successfully
developed or released as commercial varieties with yields com-
parative to other soybean varieties in the United States. Among
them are soybeans with linolenic acid as low as 3%, stearic as
high as 20%, and oleic as high as 80%. Oils from these lines are
targeted for specific applications for their functional advan-
tages over common soybean oil. Such effort will provide ben-
efits to food and industrial users and could increase the value
of soybeans as an oilseed.
Soybean Protein Products
The largest nutritionally valuable component present in soy-
beans is protein. Unlike soybean oil, which is used mainly for
human consumption, soybean protein is used largely as feed-
stuff. Only a small portion is for direct human consumption as
either traditional soyfoods or protein ingredients. Therefore,
soybean protein has been severely underutilized in terms of
human consumption.
Soybean protein products are mostly made from defatted
soy flakes, the main coproduct of modern soybean processing.
They are not consumed directly as food, but as versatile ingre-
dients used in virtually every type of food system, including
bakery, dairy, meat, breakfast cereal, beverages, infant formu-
las, and dairy and meat analogs. In these food systems, they not
only boost protein content but also provide many functional
properties, including gelling, emulsifying, water-holding, and
fat-absorbing properties. There are four major types of soybean
protein products: flour, concentrates, isolates, and textured
soybean protein.
Soy flour is one of the least-processed soybean protein
products. Many types are available in the market, including
defatted, full-fat, and low-fat, as well as enzyme-active, toasted,
and textured. Defatted soy flour has been the most common
type. It is produced by simply grinding defatted soy flakes after
modern soybean processing, containing about 50% protein.
Full-fat soy flour is made by grinding dehulled whole soy-
beans. Low-fat soy flour can be made by extrudingexpelling
soybeans, followed by milling the cake. In general, soy flour is
primarily used as an ingredient in the bakery industry.
Soybean protein concentrate is traditionally made by aque-
ous alcohol extraction of defatted soy flakes. The resulting
product has about 70% protein with the remaining portion
being mainly insoluble carbohydrates. The product may be
further processed by thermal processing and homogenization
for better functionality. Soybean protein concentrate is widely
used in the meat industry to bind water and emulsify fat and as
a key ingredient for many meat alternatives. It is also used for
protein fortification of various types of food.
Soybean protein isolate is produced by alkaline extraction
followed by precipitation at an acid pH. The process removes
both soluble and insoluble carbohydrates. The resulting prod-
uct has a protein content of 90% and is light in color and bland
in flavor. It has many functional properties, including gelation
and emulsification. As the most refined soybean protein prod-
uct, soybean protein isolate is used in processed meat, meat
analogs, soup and sauce bases, nutritional beverages, infant
formulas, and dairy replacement.
Textured soybean proteins are made by a process that
imparts a structure, such as fiber, to a proteinaceous material.
The process is known as protein texturization. It is a unique
way to make vegetable proteins palatable. Among several pro-
cesses, thermoplastic extrusion has been the most popular one
for soybean protein texturization. In a typical thermoplastic
extrusion process, dry proteinaceous materials, predominantly

defatted soy flour or soybean protein concentrate, are mixed
with water, salts, flavorings, and/or colorants and then fed into
an extruder. Under high temperature and low moisture
(<30%) conditions, the extrudate expands rapidly upon
emerging from the die. The final product, having porous tex-
ture, comes in a variety of shapes, sizes, and colors. Its use
ranges from meat extenders to meat analogs. It must be rehy-
drated with water before use.
Western Soyfoods
Traditional soyfoods, along with soybeans, were officially
introduced to the West during the nineteenth and twentieth
centuries. Since then, some of them, such as soy sauce, have
been accepted by Westerners, whereas many others have been
modified to suite local food culture and tastes. Furthermore,
many new soyfoods have been developed. The modified and
new soyfoods are known collectively as second-generation or
modern soyfoods. They can also be called Western or Western-
ized soyfoods, simply because they were developed in the West
and may look or taste like Western foods (Figure 8). Examples
include soy ice cream, soy yogurts, soy cheese, soy butter, soy
puddings, tofu spreads, soy burgers, meatless meatballs, imita-
tion bacon bits, and imitation chicken strips.
Soybean as Animal Feed
The majority of soybeans are used as animal feed mainly in the
form of soybean meal, which is the coproduct of modern
soybean processing. On a historical note, soybean meal has
been an accepted part of livestock and poultry diets in the
United States since the mid-1930s. Soybean feed production
expanded rapidly in the mid-1970s and further accelerated in
the early 1990s due to a growing demand for animal feed from
developing countries. Prohibiting the use of slaughterhouse
by-products in animal feeds and expanding aquaculture indus-
try have also fueled the demand for this high-quality protein
source.
Soybean meal is characterized by high protein content, low
crude fiber, and high palatability. It contains high amounts of
lysine, tryptophan, threonine, and isoleucine that are often
lacking in cereal grains. Because of these features, soybean
meal is the most important protein source used to feed
Figure 8 Imitation chicken strips made with soybean and other
vegetable proteins. Courtesy of Beyond Meat (www.beyondmeat.com).
THE OILSEEDS | Soybean: Overview 235
livestock, representing two-thirds of the total world output of
protein feedstuffs, which include all other major oilseed meals
and fish meal. Over half of soybean meal processed for live-
stock feed is fed to poultry, about one-quarter is fed to swine,
and the rest is used for beef cattle, dairy cattle, and pet food.
Soybean protein is increasingly found in fish food, for both
home aquariums and commercial aquaculture. Furthermore,
limited availability and increasing cost of fish meal have led to
the use of soybean meal as a dietary protein source for a variety
of cultured marine species as well.
Soybean as Industrial Materials: The New Use
While the soybean has been traditionally used as food for
human consumption and is primarily used as animal feed at
present, its versatility and chemical composition make it an
attractive material for industrial uses as well. In recent years,
the desire for biodegradable and less toxic polymers from
renewable resources instead of fossil-based chemicals has
been intensified. This has stimulated research on using soy-
bean as an industrial material. About 15% of soybean oil is
now used as feedstock for biodiesel, emulsifiers, lubricants,
plasticizers, surfactants, plastics, solvents, resins, and more.
Soybean and other vegetable oils have desirable environmental
qualities, such as being inherently biodegradable and renew-
able, having low ecotoxicity and low toxicity toward humans,
and contributing no volatile organic chemicals. For example,
soy-based engine oil emits fewer harmful emissions than
petrochemical-based oil for increased worker safety. Soy ink
is superior to petroleum-based inks, because it is not toxic,
renewable, and environmentally friendly. When soybean oil
is converted into fatty acid methyl ester through transesterifica-
tion with methanol, the product is biodiesel. Soy biodiesel is
nontoxic, renewable, and environmentally friendly.
Soybean protein products, particularly defatted soybean
meal, also have industrial applications, mostly as adhesives
and paper coatings. These applications are essentially based
on such functional properties of soybean protein products,
including adhesive strength, texture modification, film form-
ing, heat resistance, water retention, emulsification, and pro-
tective colloid properties. For example, biocomposites are
building materials made from recycled newspaper and soy
flour. They replace other products traditionally made from
wood, such as furniture, flooring, and countertops. Particle-
board, laminated plywood, and finger-jointed lumber are
made with soy-based wood adhesives. Soy-based foams are
currently being developed for use in coolers, refrigerators,
automotive interiors, and even footwear. Soybean products
are also found in many popular brands of home and commer-
cial carpet and in auto upholstery applications.
Conclusion and Outlook
Soybeans have a unique composition and an array of old and
new uses. The past four decades have provided many break-
throughs in soybean production, processing, and end use. As
research progresses and technology advances on many fronts,
soybeans are poised to become the most important source of
236 THE OILSEEDS | Soybean: Overview
protein for humans and animals and an important renewable
source of industrial materials in the twenty-first century.
Exercises for Revision
Why is soybean considered an oilseed? What differentiates
it from other oilseeds?
What features make soybean a legume?
Why has soybean become a top cash crop in the world since
the middle of the last century?
If the typical soybean oil is healthy, why do we need to
modify it through processing and/or genetic manipulation?
Why are the majority of traditional soyfoods not favored in
the West, and why do soyfoods have to be Westernized or
localized?
Why is soybean prized for its use as animal feed at the
present time?
Do you think that genetically engineered soybeans are the
future for improving soybean production and quality?
Which type of soybean uses do you think will increase in
the future: food, feed, or industrial material?
Exercises for Readers to Explore the Topic Further
Currently, the food industry is making an effort to ban or
eliminate trans fats from food. What are the best solutions
for the soybean industry to meet this challenge?
Soyfood consumption in the West has been increasing
significantly for the past 23 decades. However, compared
with other foods, the room for growth is still huge. What
efforts need to be undertaken to further increase soyfood
consumption?
Industrial use of soybeans has been considered new. What
research needs to be carried out to further increase this type
of use?
See also: Agronomy of Grain Growing: Soybean: Agronomy;
Breeding of Grains: Soybean: Germplasm, Breeding, and Genetics;
Food Grains and the Consumer: Genetically Modified Grains and
the Consumer; Food Grains and Well-being: Nutrition: Soy-Based
Foods; Grain Marketing and Grading: Soybean: Grading and
Marketing; Grains Around the World: Grain Production and
Consumption: Oilseeds in North America;Non-wheat Foods:
Soybean: Soy-Based Fermented Foods; Soybean: Soymilk, Tofu, and
Okara; Oilseed and Legume Processing: Soybean: Processing;
Soybean: Soy Concentrates and Isolates; Scientific Organizations
Related to Grains: Research Organizations of the World: CGIAR;
The Basics: Taxonomic Classification of Grain Species; The Grain
Chain: The Route from Genes to Grain-Based Products; The Grains that
Feed the World; The Legumes and Pseudocereals: Grain
Legumes and Their Dietary Impact: Overview; The Oilseeds: Oilseeds:
Overview; The Basics: The Grain Crops: An Overview.
Further Reading
Anderson JW, Johnstone BM, and Cook-Newell ML (1995) Meta-analysis of the effects
of soy protein intake on serum lipids. New England Journal of Medicine 333: 276.
Campbell KA, Glatz CE, Johnson LA, et al. (2011) Advances in aqueous extraction
processing of soybeans. Journal of the American Oil Chemists Society
88: 449465.
Chen KI, Erh MH, Su NW, Liu WH, Chou CC, and Cheng KC (2012) Soyfoods and
soybean products: From traditional use to modern applications. Applied
Microbiology and Biotechnology 96: 922.
Erickson DR (ed.) (1995) Practical Handbook of Soybean Processing and Utilization.
Champaign, IL: AOCS Press.
Isanga J and Zhang GN (2008) Soybean bioactive components and their implications to
health A review. Food Reviews International 24: 252276.
Johnson LA, White PJ, and Galloway R (eds.) (2008) Soybeans Chemistry, Production,
Processing and Utilization. Champaign, IL: AOCS Press.
Liu K (1997) Soybeans: Chemistry, Technology, and Utilization. New York: Chapman &
Hall.
Liu K (ed.) (2005) Soybeans as Functional Foods and Ingredients. Champaign, IL:
AOCS Press.
Messina M and Lane B (2007) Soy protein, soybean isoflavones and coronary heart
disease risk: Where do we stand? Future Lipidology 2: 5574.
Sikora S, Blazinkov M, Babic K, Sudaric A, and Redzepovic S (2008) Symbiotic nitrogen
fixation and sustainable soybean production. Cereal Research Communications
36: 14831486.
Stupar RM and Specht JE (2013) Insights from the soybean (Glycine max and Glycine
soja) genome: Past, present, and future. Advances in Agronomy 118: 177204.
Wang X, Cai X, and Zhang J, et al. (eds.) (1997) Zhong Guo Da Dou Zhi Ping (Chinese
Soybean Food Products) (in Chinese). Beijing: China Light Industry Publisher.
 Canola: Overview
VJ Barthet, Canadian Grain Commission, Winnipeg, MB, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Development of rapeseed into Canola
Canola production in Canada since 1943
Canola, countries exporting and importing low-erucic-acid
(LEA) rapeseed
Canola major compound contents oil, protein, and
glucosinolate
Main fatty acid present in Canola and LEA rapeseeds
Canola processing and health effects
Learning Objective
In the following article, the reader will learn about canola,
its production, composition, and processing.
Introduction
Canola is a Canadian crop and the name canola is a trademark;
it came from the contraction of Canada and ola for oil.
Canola plants belong to the Brassicaceae (Cruciferae) family
that comprises about 350 genera and 3000 species including
Brassica rapa (syn. B. campestris subsp. oleifera (turnip rape)), B.
napus (rutabagas), B. oleracea (cabbage and cauliflower), and B.
juncea (mustard). B. campestris originated in the foothills of the
Himalayan mountains; B. napus most likely originated in the
Mediterranean region and was the result of natural crosses
between B. campestris and B. oleracea plants.
Development of Low-Glucosinolate Low-Erucic-Acid
Rapeseed
Rapeseed culture was introduced in western Canada during
World War II (WWII). The seeds were crushed and the oil
was used principally as a lubricant. At the time, two types of
rapeseeds were grown: B. napus, also called Argentine canola,
and B. campestris (rapa), referred to as Polish seed. To make
lubricants, the iodine value of the oil should be in the 103105
range; it was found that B. napus was ideal, with an iodine value
close to 105, whereas Polish seeds (B. campestris) were not as
suitable since the iodine value of the oil was close to 115. A
breeding program was begun to develop rapeseed varieties
with improved agronomy such as uniformity and lodging
and an increased oil content while lowering the iodine value
of the oil.
Later on, rapeseed almost disappeared from the prairies
because the oil demands changed and other oilseed crops
were available. However, rapeseed had shown that its culture
was quite favorable for western Canada since it needed fewer
heat units to grow than the other oilseeds (soybean or
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00029-2
sunflower). Oil uses became easier since work on rapeseed oil
processing, refining, bleaching, and deodorizing showed that
rapeseed oil could be used as an edible oil, as soybean oil was.
This led to the use of rapeseed oil as an edible oil in Canada.
However, work done in the early 1950s by Caroll on rats
showed that rapeseed oil, and erucic acid in particular, had a
negative effect on the heath of the rats. This led, in 1956, the
Food and Drug Directorate of the Department of National
Health and Welfare (now Health Canada) to prohibit rapeseed
oil in human food. Even though this directive was canceled
within months, it gave a very strong push to the Canadian
rapeseed breeding programs to develop a low-erucic-acid
(LEA) rapeseed. These breeding programs were located in the
prairies, Winnipeg with Dr. Stefansson (University of Mani-
toba) and Saskatoon with Dr. Downey (Dominion Forage
Crop Laboratory, now Agriculture and Agri-Food Canada).
In 1958, a first variety trial was conducted with six varieties
(Argentine, Arlo, Golden, Gute, Polish, and Regina II) of rape-
seed (B. napus and B. campestris) grown in 22 locations
throughout the western provinces. This led in 1961 to the
first paper showing an effect of variety and environment on
the fatty acid composition of the oil for all fatty acids except
linolenic acid. The paper also showed an inverse relationship
between oleic acid content and erucic acid content for both B.
napus and B. campestris. The author concluded by saying sin-
gles plant selection and crosses should yield valuable informa-
tion of individual fatty acid and the development of gas liquid
chromatography (GC) makes it possible to do such studies.
Truly, the development of LEA rapeseed became linked to the
development of GC, since precise and short-time analysis was a
requirement to analyze accurately a large number of samples.
To look for LEA rapeseed (and high), single-plant analyses
of Liho (a variety of B. napus from Germany), known for its LEA
content compared to other varieties, were conducted and large
variations in erucic acid content were observed (650%). This
led to the first LEA rapeseed plants; progeny of selected LEA
plants, grown in both Winnipeg and Saskatoon, gave plants
with seeds containing 0% erucic acid in their oil in both
locations. In 1963, Downey and Harvey developed a technique
where only half of the seed was analyzed for the fatty acid
composition while the other half, formed by half of the coty-
ledon attached to the radicle, was germinated and planted. This
innovative technique allowed growing plants with a better
knowledge and control of the oil composition. Subsequent
work showed that the fatty acid composition was controlled
by the genotype of the developing seeds and not by the mater-
nal plant, and several genes were involved and their actions
were additive. Later on, it was shown that five genes (e, Ea, Eb,
Ec, and Ed) controlled the level of erucic in B. napus in an
additive manner.
Work was also conducted with B. campestris (rapa) to get
LEA B. campestris varieties. In 1964, Downey reported on the
selection of inbred seed from a Polish LEA variety of
237
238 THE OILSEEDS | Canola: Overview
B. campestris having no erucic acid in its oil. Then, Dorrel and
Downey showed that fatty acid composition (erucic acid con-
tent) of B. campestris was as in B. napus, controlled by the
genotype of the developing seed; however, they could not
confirm their hypothesis that a single gene controls the erucic
level. Instead, they showed that the fatty acid composition was
affected by the stage of the seed development.
Simultaneously, work was started to reduce the glucosino-
late content of the seeds since several studies had shown that
glucosinolates had a negative effect on animal health. All these
nutritional studies were summarized in 1965 by Bowland et al.
in a report. In 1959, the first Canadian report showing signif-
icant variations of isothiocyanate contents between varieties
but not locations was published. Important differences were
reported between Argentine-type (B. napus L.) rapeseed and
Polish-type (B. campestris (rapa) L.) rapeseed. In 1970, Kondra
and Downey reported that, opposite to erucic acid, glucona-
pin, glucobrassicanapin, and progoitrin (the main glucosino-
lates) contents were determined by the genotype of the
maternal plant. The work was done using Bronowski, a variety
from Poland containing low levels of the three glucosinolates,
and Target, a Canadian variety with high levels of glucosino-
lates. Work was also done in Europe with winter rape to
develop low-glucosinolate rapeseeds also using the Bronowski
variety.
This led in 1974 to the registration of Tower, the first
summer rape (B. napus) with LEA content in the oil and low-
glucosinolate content of the meal. In 1977, the first B. campes-
tris low in erucic acid and in total glucosinolates, called Candle,
was registered in Canada by the Agriculture and Agri-Food
Canada Saskatoon Research Centre.
In 1978, the Rapeseed Association of Canada decided to
name this new crop (low-glucosinolate LEA or double-low
rapeseed) canola, a name made from the condensation of
can from Canada to represent the Canadian origin of the
new crop and ola to follow some famous brand name exam-
ples (cola, Mazola, etc.). Canola is now defined as a rapeseed
that produces an oil that must contain less than 2% erucic acid,
and the solid component of the seed must contain less than
30 mmol of any one or any mixture of 3-butenyl glucosinolate,
4-pentenyl glucosinolate, 2-hydroxy-3-butenyl glucosinolate,
and 2-hydroxy-4-pentenyl glucosinolate per gram of air-dry,
oil-free solid (http://www.canolacouncil.org/oil-and-meal/
what-is-canola/).
Canola varieties can belong to any of the following three
species: B. napus L., B. rapa (campestris) L. (with the three
subspecies B. rapa subsp. oleifera, B. rapa subsp. trilocularis,
and B. rapa subsp. dichotoma), and B. juncea L. Czern. However,
in Canada, most canola varieties grown belong to the B. napus
L. species. In Europe, canola is referred as low-glucosinolate
LEA rapeseed.
The work to produce LEA varieties showed how high-erucic-
acid rapeseed (HEAR) varieties could also be produced. This
led to the development of rapeseed with very high levels of
erucic acid so the oil had applications in the lubricant industry
and with very low levels of glucosinolates so the meal could be
used by the feed industry. One of the first varieties with very
high level of erucic acid and low glucosinolate in the meal was
Hero 50.2% erucic acid in the oil and 15 mmol g
1 of total
glucosinolate in the meal. Due to their high levels of erucic
acid, these varieties are not canola varieties; they are low-
glucosinolate HEAR. An important breeding program to
develop varieties with very high erucic acid and low glucosino-
lates is still present at the University of Manitoba.
In order to increase variation and create new potential for
breeding, ethyl methanesulfonate was tested on B. napus to
promote mutations. As a result, in 1987, Stellar was registered.
It was the first low-linolenic (3.3%), high-linoleic (28.9%)
canola (0.1% erucic acid in the oil and 8 mmol g
1 glucosino-
late in the meal) variety registered in Canada.
Production
Rapeseed has been an important source of edible vegetable oil
in Asia for almost 4000 years, used as a lighting oil and edible
oil in Europe since at least the middle ages. However, in
Canada, rapeseed production started during WWII as a source
of high-quality lubricant for marine engines. It was a very small
production about 1000 metric tonnes in 1943 (Figure 1). The
development of the first canola variety in 1974 led to an
increase in production and a switch from rapeseed to canola.
By 1977, almost 2 million tonnes of canola was produced in
western Canada. Canola production increased steadily up to
1995; then, the introduction of the first herbicide-resistant
varieties led to another jump in production (Figure 1). This
was followed by the development of new varieties, mostly
hybrid, with better agronomic traits allowing a rapid produc-
tion increase since 2000. Starting in 2004, the demand in
canola increased due to a sharp increase in the world biodiesel
production. This demand was matched by an increase in pro-
duction due to varieties with better yields. In 2013, western
Canadian canola production reached 17.96 million tonnes
(Figure 1). Canola is considered as the Cinderella crop in
Canada, from a forgotten crop to a major cash crop for Cana-
dian farmers. Today, almost all canola varieties grown in Can-
ada are herbicide-tolerant. Three types are found, two were
obtained by insertion of a gene and are GMO (Roundup
18,000
16,000
Production (metric tonnes 1000)
14,000
12,000
10,000
8000
6000
4000
2000
0
1940 1950 1960 1970 1980 1990 2000 2010
Year
Figure 1 Canadian production of rapeseed and canola in metric tonnes
(1000) from 1943 to 2013.
 THE OILSEEDS | Canola: Overview 239

Ready and LibertyLink), and one was obtained by mutation
and is a non-GMO (Clearfield).
The world canola and LEA rapeseed production is con-
centrated in the European Community, Canada, and Aus-
tralia (Table 1). China and India are important rapeseed
producers; however, most of the rapeseed crops grown in
these countries are not low in erucic acid and low in glu-
cosinolate (Table 3).
Canola/LEA rapeseed trade data are presented in Table 2.
The European Community, China, Japan, and Mexico are the
main importing countries, whereas Canada, Australia, and
Ukraine are the main, if not the only, exporting countries.
Canada is the main canola exporter; since 2004, Canada has
been responsible of 70% of the world total canola exports. The
United States is the main canola customer for Canada, import-
ing an equivalent of 5 million tonnes of seeds; however, less
than 0.5 million tonnes of seeds is exported to the United
States from Canada. Canadian canola export to the United
States is in the form of oil, about 1.5 million tonnes of canola
oil. There is also an important export in the form of canola
meal well over 2.5 million tonnes mainly used by the US
dairy industry.
Growing Conditions
Canola, in Canada, is a summer crop. It is seeded in spring
(May) and harvested from August to October. In Europe and
China, LEA rapeseed and rapeseed are winter crops; they are
seeded in August and harvested the next year from early July to
August. Australia also grows canola, but opposite to Canada,
Australian canola is a winter crop seeded at the end of
summerearly autumn and harvested the following year end
of springearly summer.
Composition
Canola is marketed for its oil. Therefore, canola quality can be
defined by its oil content and the fatty acid composition of the
oil. Canola meal (what is left of the seed after crushing) is used
for animal feed so the glucosinolate content is also an impor-
tant parameter in determining canola seed quality for feed.
Table 3 presents the proximate analysis and the relative fatty
acid composition of canola and rapeseed grown in various part
of the world.
Processing and Uses
Canola seeds (LEA rapeseed and rapeseed) are processed by
crushing plant to produce oil and meal. When the seed is
delivered to the crushing plant, they are stored in bins to be
processed. Processing steps might depend on the crushing plan
and the country. In Canada, the weather is usually dry and
cold, so the first step of the processing is to pass the seed
through a conditioner with heat and moisture allowing better
flaking and to heat the seed, making the oil more fluid and

Table 1 Top five canola/rapeseed producing countries and world production for the last 10 years (1000 tonnes)

Country

Year European Union Canada China India Australia World total

200304 11 214 6771 11 420 6800 1703 39 464

200405 15 458 7673 13 182 6500 1542 46 144
200506 15 564 9483 13 052 7000 1419 48 589
200607 16 112 9000 10 966 5800 573 45 155
200708 18 397 9611 10 573 5450 1214 48 560
200809 19 062 12 644 12 100 6700 1844 57 971
200910 21 633 12 898 13 500 6400 1907 60 905
201011 20 300 12 789 12 900 7100 2359 59 891
201112 18 954 14 608 13 000 6200 3427 60 840
201213 19 300 13 310 12 900 6900 3898 61 626

Table 2 Rapeseed (canola) trade from 2004 to 2012 (1000 tonnes)

Importing country Exporting country

Year European Community China Japan Mexico Pakistan World total Canada Australia Ukraine World total

200405 224 362 2277 1196 672 5507 3756 927 160 5507
200506 712 713 2333 1259 820 7521 5846 827 357 7521
200607 433 954 2170 1065 806 7106 5426 230 755 7106
200708 1388 855 2210 1317 535 8648 5916 508 1496 8648
200809 2897 3085 2061 1077 596 11 584 7660 1111 2157 11 584
200910 2284 2059 2287 1403 940 11 146 7315 1172 1910 11 146
201011 2829 1100 2337 1492 811 10 484 7249 1507 1116 10 484
201112 3485 2487 2391 1653 932 13 071 8680 2486 1341 13 071
240 THE OILSEEDS | Canola: Overview
Table 3 Oil, protein, and total glucosinolate content ranges (dry basis) and relative fatty acid composition of canola/LEA rapeseed and rapeseed
grown in various parts of the world
Canadaa (200413)
Proximate composition (dry basis)
Oil content (%) 48.4
Protein (%) 22.5
Meal protein 43.6
Glucosinolate (mmol g
1) 12.2
Fatty acid composition (% in the oil)
Oleic acid 61.8
Linoleic acid 19.1
a-Linolenic acid 9.9
Erucic acid 0.03
a
Canadian Grain Commission data.
b
Australian Oilseeds Federation data.
c
Centre Technique Interprofessionnel des Oleagineux et du Chanvre data.
d
Li et al. (2011).
easier to extract. The second step is flaking where the seeds are
placed into big rollers in order to break the seed structure to
allow an easier crush and facilitate the oil extraction. A cooking
step follows to ease the oil extraction and to inactivate endog-
enous enzymes of the seeds. Canola contains low levels of
glucosinolates; however, it is important to ensure that they
are not hydrolyzed during the processing to guarantee good
quality for the meal. The myrosinase responsible for the glu-
cosinolate hydrolysis should be inactivated. Enzymes (lipases)
able to degrade the triglycerides must also be inactivated to
ensure no oil degradation occurs during the processing. Then,
the seeds are submitted to a screw press (expeller press) that
squeezes the oil out of the seeds. Not all oil is recovered at this
step. A solvent extraction, usually with hexane, is performed on
the expeller press cake. After solvent extraction, depending on
the crushing plan, up to 99% of the oil could be recovered
from the seeds. The last step of canola processing is a refining
process during which undesirable substances are removed
from the crude oil to give the yellowish oil found in
supermarkets.
Chlorophyll is present in canola seeds when the seeds are
harvested before reaching full maturity. Canadian canola seeds
usually contain higher levels of chlorophyll because the grow-
ing season is very short May to September compared to
canola grown in Europe, Australia, or China, September to
June. Chlorophyll is an undesirable substance in canola oil
and has to be removed. Chlorophyll with its heme structure
and the magnesium ion in the center is highly reactive and can
oxidize unsaturated fatty acids in the oil. Canola oil contains
about 20% linoleic acid (C18:2) and 10% a-linolenic acid
(ALA) (C18:3); these fatty acids can oxidize 10 (linoleic acid)
and 50 times (ALA) faster than oleic acid (C18:1). To protect
the canola oil from oxidation and to allow further processing
of the oil, chlorophylls have to be removed during refining.
Refined canola oil can be further processed to give other
food products such as margarines and shortenings or industrial
products such as biodiesel.
Canola meal, the leftover from the seed once the oil is
removed, can also be further processed. It is then used by the
feed industry as pellet or mash product. Canola meal is con-
sidered a premium ingredient in dairy diets where it has been
Australiab (200012) Francec (200413) Chinad (200010)
44.546.8 43.6c 39.742.6
23.625.3
40.246.4 38.8c
4.310.6 14.5 27.638.0
59.063.0 62.9d
18.220.4 19.6d
9.011.0 9.3c
<0.1 3.1416.3
shown to increase milk production when compared to diets
containing cottonseed or soybean. It is also used in poultry and
swine feeds.
Food Uses and Health Effects
Canola oil is considered a very healthy oil due to its fatty acid
composition. It averages about 60% oleic acid (C18:1), 20%
linoleic acid (C18:2), and 10% ALA (C18:3) (Table 1). This
makes canola oil a good source of ALA with the ideal ratio
(2:1) of omega-6 (o-6) to omega-3 (o-3). When compared to
the other vegetable oils, it has the lowest amount of saturated
fatty acid (7% or less) compared to sunflower (12%). Studies
with diets containing canola oil suggested that canola could
reduce blood cholesterol levels and reduce the risk of cardio-
vascular disease. Canola oil can be used by food manufactures
to produce various kinds of salad dressing. It is also found at
local food stores as a salad oil, which can also be used as
cooking oil. Margarines with canola can also be found at
food markets and used by processors for baking.
In response to the mandatory labeling of trans-fatty acids in
food products, canola varieties with high-stability oils
(ALA < 3%) were grown. These new oils have higher levels of
oleic acid (6574% vs. 60%) and lower levels of ALA (13% vs.
10%) than the conventional canola oil. These high-stability
oils are used by the food industry for frying, allowing the
reduction/elimination of trans-fatty acids from fried foods
(e.g., potato chips) making these products a healthier choice.
In North America, some chip brands and other fried snacks are
processed using high-stability canola oil.
Exercises for Revision
What are the three Brassica species that can be called canola?
What are the major producing and importing countries for
canola seeds?
What is the main fatty acid in canola oil?
What were the two compounds reduced from rapeseed to
give canola?

What is the ratio of omega-6/omega-3 for conventional
canola?
Why must chlorophyll be removed from canola crude oil?
Exercises for Readers to Explore the Topic Further
What are the advantages of modifying the fatty acid com-
position of canola seeds?
Why can changes in weather patterns affect the quality
parameters (chlorophyll, oil, and fatty acid composition)
of Canadian canola seeds?
See also: Breeding of Grains: Canola/Rapeseed: Genetics and
Breeding; Food Grains and the Consumer: Genetically Modified
Grains and the Consumer; Grain Harvest, Storage and Transport:
Canola: Harvest, Transport, and Storage; Grains Around the World:
Grain Production and Consumption: Oilseeds in North America;
Oilseed and Legume Processing: Canola: Processing; The
Basics: The Grain Crops: An Overview.
Further Reading
Abraham V and de Man JM (1986) Hydrogenation of canola oil as affected by
chlorophyll. Journal of the American Oil Chemists Society 63(9): 11851188.
Anand IJ and Downey RK (1981) A study of erucic acid alleles in digenomic rapeseeds
(Brassica napus L.). Canadian Journal of Plant Science 61: 199203.
Aukema H and Campbell L (2011) Oil nutrition and utilization. In: Daun JK, Eskin NAM,
and Hickling D (eds.) Canola, Chemistry, Production, Processing and Utilization,
pp. 245280. Urbana, IL: AOCS Press, Chapter 9.
Bowland JP, Clandinin DR, and Wetter LR (1965) Oil and Meal from Canadian
Rapeseed. Ottawa, Canada: Department of Agriculture, Publication # 1257. https://
archive.org/details/oilmealfromcanad00bell.
Busch H and Robbelen G (1981) Neidriger glucosinolatgehalt als zuchtzeil fu
winterraps. Agew. Botanik 55: 361371.
Craig BM (1961) Varietal and environmental effects on rapeseed III. Fatty acid
composition of 1958 varietal test. Canadian Journal of Plant Science 41: 204210.
Downey RK and Harvey BL (1963) Methods of breeding for oil quality in rape. Canadian
Journal of Plant Science 43: 271275.
Fowler DB and Stefansson BR (1972) Effects of the mutagenic agent ethyl
methanesulfonate on the M1 generation of rape (Brassica napus). Canadian Journal
of Plant Science 52(1): 5362.
Fowler DB and Stefansson BR (1975) Ethyl methanesulfonate-induced mutations in
rape (Brassica napus). Canadian Journal of Plant Science 55(3): 817821.
Harvey BL and Downey RK (1964) The inheritance of erucic acid content in rapeseed
(Brassica napus). Canadian Journal of Plant Science 44: 104111.
THE OILSEEDS | Canola: Overview 241
Jahries G and Schafer U (2011) Rapeseed (Brassica napus) oil and its benefits for
human health. In: Preedy V, Watson R, and Patel V (eds.) Nuts and Seeds in Health
and Disease Prevention, 1st ed., Chapter 114, pp. 967974. ISBN 9780123756893.
Krzymanski J and Downey RK (1969) Inheritance of fatty acid composition in winter
forms of rapeseed Brassica napus. Canadian Journal of Plant Science 49: 313319.
Le MHA, Landero JL, Beltranena E, and Zijlstra RT (2014) The effect of feeding
increasing inclusion of extruded Brassica juncea canola expeller on growth
performance and nutrient digestibility in weaned pigs. Animal Feed Science and
Technology 192: 7380. http://dx.doi.org/10.1016/j.anifeedsci.2014.02.006.
Li P, Ding X, and Zhou H (2011) Rapeseed quality improvement and monitoring plan in
China from 2000 to 2010. In: Proceeding of the 13th International Rapeseed
Congress, Prague, Czech Republic, 59 June 2011.
Mag TK (1983) Canola oil processing in Canada. Journal of the American Oil Chemists
Society 60(2): 380384.
Martineau R, Ouellet DR, and Lapierre H (2013) Feeding canola meal to dairy cows: A
meta-analysis on lactational responses. Journal of Dairy Science 96(3):
17011714.
Mulrooney CN, Schingoethe DJ, Kalscheur KF, and Hippen AR (2009) Canola meal
replacing distillers grains with solubles for lactating dairy cows. Journal of Dairy
Science 92(11): 56695676.
Scarth R, McVetty PBE, Rimmer SR, and Stefansson BR (1988) Stellar low linolenic-
high linoleic acid summer rape. Canadian Journal of Plant Science 68: 509511.
Scarth R, McVetty PBE, Rimmer SR, and Stefansson BR (1991) Hero summer rape.
Canadian Journal of Plant Science 71: 865866.
Slominski BA, Simbaya J, Campbell LD, Rakow G, and Guenter W (1999) Nutritive value
for broilers of meals derived from newly developed varieties of yellow-seeded
canola. Animal Feed Science and Technology 78(34): 249262.
Stefansson BR and Hougen FW (1964) Selection of rape plants (Brassica napus) with
seed oil practically free of erucic acid. Canadian Journal of Plant Science 44(4):
359364.
Stefansson BR, Hougen FW, and Downey RK (1961) Note on the isolation of rape plants
with seed oil free from erucic acid. Canadian Journal of Plant Science 41(1):
218219.
Stefansson RR and Kondra ZP (1975) Tower summer rape. Canadian Journal of Plant
Science 55: 343344.
Stricker H, Duchini F, Facchini M, and Mombelli G (2008) Canola oil decreases
cholesterol and improves endothelial function in patients with peripheral arterial
occlusive disease A pilot study. Artery Research 2(2): 6773.
Tautorus CL and Low NH (1993) Chemical aspects of chlorophyll breakdown products
and their relevance to canola oil stability. Journal of the American Oil Chemists
Society 70(9): 843847.
Relevant Websites
http://www.australianoilseeds.com/oilseeds_industry/quality_of_australian_canola
Australian Oilseeds Federation.
http://www.cetiom.fr/index.php?id15446 Centre Technique Interprofessionnel des
Oleagineux et du Chanvre.
http://grainscanada.gc.ca/canola/hqcm-mqrc-eng.htm Canadian Grain Commission.
http://www5.statcan.gc.ca/cansim/a26?langeng&retrLangeng&id0010010&
paSer&pattern&stByVal1&p11&p231&tabModedataTable&csid
Statistics Canada.
 Cottonseed: Overview
E Hernandez, Texas A&M University, College Station, TX, USA
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 1, pp 343348, 2004, Elsevier Ltd.
Topic Highlights
Traditional varieties of cottonseed contain gossypol
(0.62.0% in raw cottonseed kernels).
Gossypol is a polyphenolic compound that is toxic to man
and monogastric animals, presumably functioning as a
natural insecticide for the plant.
Gossypol is located in glands, appearing as black specks
in the stems, leaves, and seed.
The breeding of glandless cotton varieties (since the
1960s) means that cottonseed is available without gossypol.
Meal and oil from glandless cottonseed are similar in com-
position to these products from the traditional glanded
seeds.
Glandless cottonseed offers the advantages for processing
of reduced energy costs, elimination of prepress operations,
and the absence of color in the resulting oil.
Food products from glandless cottonseed include whole
roasted seeds, flour, protein concentrates, and isolates.
Glandless cottonseed is suitable to feed monogastric ani-
mals, not only due to the absence of gossypol but also
because of higher available energy.
Learning Objectives
Understand the important differences between glandless
and the traditional glanded cotton varieties.
Become familiar with the processing involved in producing
the various products from cottonseed for food and feed
uses.
Introduction
Most varieties of cotton (Gossypium hirsutum L., G. aboreum L.,
G. barbadense L., or G. herbaceum L.) are grown mainly in warm
climates around the world. Over 98.5 million tonnes (Mt) of
cotton were produced worldwide in 200102 and over 33.6 Mt
of cottonseed. Cotton is grown for its fiber (over 80% of its
value) and the seed is used mostly for oil recovery and feed.
Whole cottonseed can be fed to dairy cattle, and the meal
resulting from oil extraction is fed primarily to ruminants
and, in limited amounts, to poultry and swine.
Traditional varieties of cottonseed contain gossypol, a
yellow-green polyphenolic compound considered toxic to
man and monogastric animals (Figure 1), reportedly affecting
the heart, liver, and reproductive organs. It has been used in
China as a male contraceptive but the practice was abandoned
because of permanent side effects. Gossypol is dispersed in
the plant as deposited structures or glands, which can be
seen as black specks in the stems, leaves, and seed (Figure 2).
242 Encyclopedia of Food Grains, Second Edition
The glands in the seed are ovoid structures containing 3550%
gossypol and are 0.0250.178 mm in diameter. These gossypol
glands are difficult to break by mechanical means, but
heat generated in extraction of oil by pressing, binds gossypol
to protein, turning it nontoxic. Switching to solvent extraction
with hexane, where no appreciable heat is generated, increases
the free gossypol content in the meal over ten times.
Raw cottonseed kernels may contain 0.62.0% free gossy-
pol. The Food and Drug Administration in the US (FDA) limits
free gossypol in human food products and ingredients at
450 ppm, and the Protein Advisory Group of the United
Nations Food and Agriculture Organization (FAO) and
World Health Organization (WHO) has set maximum guide-
lines of 600 ppm for free gossypol and 12 000 ppm total gos-
sypol. The feed industry in the US set limits for free gossypol
levels in poultry diets at 100 ppm maximum for broilers and
40 ppm for laying hens.
Removal by mechanical separation of the gossypol-contai-
ning glands has also been reported. Also addition of iron
salts, such as ferrous sulfate, which bind the gossypol in
feeds and render it biologically inactive, is practiced in some
countries.
Solvent extraction with hexane of cottonseed oil, com-
monly used commercially, removes only small amounts of
gossypol with the oil. However, extraction with more polar
solvents is effective in removing gossypol. Examples of polar
solvents used include aqueous acetone, a mixture of acetone
and hexane, and sequential extraction with hexane, aqueous
acetone, and anhydrous acetone. Blends of methylene chloride
and hexane-acetic acid have also been used. These solvents are
not widely used commercially.
Techniques have also been reported that mill cottonseed in
the presence of hexane and then remove the intact, heavier
gossypol glands by the liquid cyclone process. Also, an air
classification process has been developed to separate intact
gossypol glands from solvent-extracted ground flour.
Glandless Cottonseed
Some of the first studies on the development of glandless
cottonseed in the US were done in the early 1950s. These
studies described that selection of plants from the Hopi
Moencopi variety could result in almost complete elimination
of pigment glands from leaves and bolls. When crossing Moen-
copi with upland cotton varieties (G. hirsutum), it was found
that glandless seed appeared in later segregating generations.
Production of glandless cotton was first conducted in Iguala,
Mexico in 1960.
Basically two genes, gl-2 and gl-3, in the plant are found to
control the production of gossypol pigments in the seed. When
present in the homozygous recessive condition (gl2gl2gl3gl3),
http://dx.doi.org/10.1016/B978-0-12-394437-5.00025-5

Figure 1 Gossypol.
Figure 2 Cross-section of glanded cottonseed (left) and glandless
seed (right). (Data from Miravalle RJ (1972) J. Am. Oil Chem. Soc. 49:
2426.)
all parts of the plant aboveground, including the seed, have
no pigment glands. The presence of pigmented glands can
be easily seen as black specks by cross-sectioning a seed
(Figure 2).
Recently, the value of the whole seed as cattle feed has
increased making it less profitable to extract the oil and sell
the meal. In some cases cottonseed is left at the gin in exchange
for ginning and baling services. Sometimes glandless cotton-
seed is considered to produce fiber of lower yield and quality
than traditional glanded varieties. Some studies have shown
that the glandless factor in itself does not decrease fiber yield or
quality as long as care is taken to prevent weed and insect
infestation.
Gossypol is regarded also to be a natural insecticide. Reports
show that glandless cotton attracts more insects than glanded
varieties. Entomology studies have shown that cotton bollworm
(Heliothis zea Boddie), tobacco budworm (H. virescens F.), pink
bollworm (Pectinophua gossyprella Saunders), and lygus insects
prefer glandless cotton. It is generally recognized that glandless
varieties require closer supervision to intercept and control
insect infestations.
Processing and utilization of glandless cottonseed was
reported in the early 1960s in the US. Glandless cottonseed
oil and meal were evaluated in poultry broiler and layer
rations. Glandless cottonseed meal was found nearly as effec-
tive as soybean meal in achieving broiler gains and did not
THE OILSEEDS | Cottonseed: Overview 243
produce the green yolk discoloration in eggs commonly expe-
rienced with feeding glanded cottonseed meal.
The first commercial glandless cotton variety was a storm-
proof, boll-type cotton. It was evaluated in farm-scale trials and
made available for commercial planting in the late 1960s in the
US. Commercial sale of glandless cottonseed began in the early
1970s, with FDA approval. In 1976, under title 21 (Food and
Drugs) of the Code of Federal Regulations, sale of glandless
cottonseed kernels and cottonseed flour as food additives was
allowed. It included restrictions on gossypol (450 ppm). In
1978 the National Cottonseed Products Association (NCPA)
in the US established grades of glandless cottonseed products:
class A, to contain not more than 400 ppm of total gossypol;
class AA, to contain not more than 100 ppm total gossypol;
and class AAA, to contain not more than 10 ppm total gossy-
pol. Glandless cottonseed kernels started to be sold for use as
snack foods, in baked goods and in soft candy.
Research and production of glandless cotton for food and
feed is also reported in Europe, Africa, and Asia. Some US
cotton varieties were adapted to African crops. Development
and study of some Egyptian varieties of glandless cotton have
also been reported. From studies done in the Ivory Coast, it has
been suggested that glandless cottonseed cultivation and oil
mill processing is more economically viable in that part of
Africa. Conditions there may be better for glandless cotton
considering that in larger industrialized countries other com-
petitive oil seed proteins are more readily available. An addi-
tional advantage is that large, isolated land areas can be
dedicated to this crop and less problems of cross-pollination
would be encountered.
Processing and Products
Other than gossypol content very few differences are found in
glandless compared with glanded cottonseeds. Evaluations of
eight varieties each of glanded and glandless cottonseed were
reported (Table 1). Amino acid profiles were similar for
glanded and glandless seed. The storage and processing char-
acteristics of glandless and glanded cottonseed are also
reported to be essentially identical with regard to development
of free fatty acids, moisture levels, and refining losses in the
crude oils.
No major differences were encountered either in solvent
extraction of glanded versus glandless cottonseed. Extraction
rates of oil from glandless cottonseed flakes are about the same
as from glanded flakes, using commercial hexane. It has also
been suggested that glandless cottonseed can also be a source
of a new type of lecithin. Cottonseed has the highest content of
phospholipids after soybeans, present at about 2.2% in the oil,
but due to gossypol presence it was not commercially viable.
Currently, the major commercial source of lecithin is soybean
oil. Composition of cottonseed phospholipids consists of
phosphatidylcholine, phosphatidylethanolamine, and phos-
phatidylinositiol (33%, 22%, and 37%, respectively) and is
considered more oxidatively stable than soybean phospho-
lipids due to lower content of unsaturated fatty acids.
One of the main advantages of milling glandless cottonseed
is that it simplifies direct solvent extraction. Current commercial
practice of refining glanded cottonseed oil requires that the oil
244 THE OILSEEDS | Cottonseed: Overview

Table 1 Comparative analysis means of eight varieties each Advantages of processing glandless cottonseed over glanded
of glanded and glandless cottonseed and their productsa seed include:

Glanded Glandless 1. reduction in electrical energy used for flaking;

Product and assay cottonseed cottonseed 2. elimination of prepress operations with larger solvent
extractors with lower maintenance costs, and reduction in
Whole cottonseed
processing energy;
Oil (%) 21.0 21.1
3. since there is no color setting problem, crude cottonseed oil
Iodine no. 108.9 109.9
Protein (N  6.25; %) 23.1 22.5 can be held longer in storage;
Wt. 100 fumed kernels (g) 10.0 10.6 4. elimination of more dangerous miscella (oil-solvent solu-
No. fumed seed/100 ml 542 510 tion) processing or on-site conventional refineries;
% Kernels in lint-free seed 61.7 59.6 5. lower refining loss of glandless seed oil due to reduced use
Cottonseed kernels of alkali to eliminate color;
Oil (%) 37.8 39.7 6. reduction of bleaching earth needs; and
Protein (N  6.25; %) 39.3 38.9 7. marketing of a light-colored, gossypol-free oil.
Crude fiber (%) 1.6 1.7
Total phosphorus (%) 0.8 0.9 Some emulsification problems have been reported in refining
Total sugars (%) 7.4 6.8 desolventized crude due to higher concentration of lecithin.
Total gossypol (%) 1.2 0.02 There are some drawbacks when switching to glandless
Wt. 100 kernels (g) 6.5 7.0 crops; varying levels of gossypol can exist among individual
No. kernels/100 ml 912 844 seeds from the same plant. It is also difficult growing large
Hexane-extracted flour (meal)
acreages of glandless cottonseed, without cross-fertilization
Oil (%) 0.8 0.8
by windblown or insect-carried pollen of glanded varieties. In
Protein (N  6.25; %) 63.2 62.6
Crude fiber (%) 2.7 2.8
order to preserve the variety and seed lot purity in the produc-
Ash (%) 8.0 7.8 tion of glandless cottonseed products, it is important to mon-
Total phosphorus (%) 1.3 1.4 itor both, cross-fertilization from cotton plants in nearby fields
Total sugars (%) 13.4 13.7 and reversion to the glanded condition during successive
Total gossypol (%) 1.6 0.02 plantings due to fertilization between heterozygous plants.
Color, Hunter L values Furthermore, segregated handling of glandless cottonseed in
Dry 84.3 89.8 gins and oil mills can also be costlier. As a result large cotton-
Wet (5 water:1 flour) 48.1 71.3 seed processors may be hesitant to handle glandless cottonseed
Crude oil
and dedicate separate processing facilities.
Cyclopropenoid fatty acids (%) 0.23 0.23
Fatty acids (%)
Myristic 0.9 0.7
Palmitic 23.0 22.6
Stearic 2.2 2.1 Foods Uses
Oleic 17.7 17.7
Linoleic 55.8 56.5 Several food uses of glandless cottonseed kernels have been
Unknown 0.4 0.4 reported as well as applications on the use of glandless cotton-
Refined oil seed blended with other foods. Even though glanded cotton-
Refined oil color, red 6.9 3.7 seed is not considered suitable for human consumption, there
Bleached oil color, red 2.9 2.2 have been cases where cottonseed flour with bound gossypol
a
Dry weight basis.
has been used in nutrition intervention feeding products, such
(Source: Lawhon JT, Cater CM, and Mattil KF (1977) Journal of the American Oil as Incaparina in South America.
Chemists Society 54: 75.) Food products from glandless cottonseed include whole
roasted seeds, flour, protein concentrates, and isolates. From
these products many foods were reported to be prepared. The
be refined in its miscella state (oilhexane solution) right after technologies for the production of high-protein products have
extraction. This is necessary because if the solvent is removed been well established and apply similarly to all vegetable pro-
from the oil prior to refining, the color fixes into the oil and is teins. Specific technologies were developed for the production
very difficult to remove in subsequent bleaching operations. of cottonseed protein products that were later applied for
This means the equipment in refining of miscella has to be processing of other vegetable proteins like soybean.
explosion-proof, which is more expensive and more difficult to The preliminary steps in preparing glandless cottonseed
maintain. This is no longer the case with glandless seed proces- products are similar to the process used with glanded cotton-
sing, where binding of gossypol (by cooking and hard press or seed, i.e., ginned cottonseed is first cleaned to remove dirt and
prepress) or miscella refining become unnecessary. The oil from other impurities, conditioned, de-hulled, and separated to pro-
glandless seed is in lighter colored oil and the meal has greater duce kernels. The kernels may be size-sorted and the larger
amounts of soluble protein. Protein solubility of processed particles roasted and color-sorted. The whole kernels are sepa-
glandless cottonseed flakes or press cake has been reported to rated for roasting or conditioning. The smaller or broken
be higher than that of glanded seed (89.6%) compared with kernels may then be further conditioned, flaked, and solvent-
84.8% for extracted flakes, and 59.3% for press cake. extracted for production of flours, concentrates, and isolates.

Whole glandless cottonseed kernels are prepared either for
direct use as whole nut substitutes or for food ingredients.
These are also used in the production of high-quality flours
and food protein concentrates or isolates. The use of glandless
cottonseed kernels for the production of nutlike products by
means of various roasting methods has been reported, includ-
ing dry roasting at several temperatures, roasting under vac-
uum followed by steam injection, pressure steaming followed
by oven roasting, and deep fat frying in various oils (including
corn, cottonseed, peanut, safflower, soybean, and sunflower
oils). When the main product is to produce high-protein
flours, less care needs to be taken to minimize breakage during
de-hulling.
Glandless cottonseed flour can be prepared by extracting de-
hulled glandless seed flakes with hexane. The extracted flakes are
then desolventized and ground to a desired mesh size. In the
production of cottonseed concentrates, the defatted flakes are re-
extracted with acidified water or ethanol to remove the soluble
sugars and flavor compounds, and dried to produce protein
concentrate. A spray dryer can also be used in preparing con-
centrates where the defatted flakes are ground prior to extrac-
tion. Concentrates have been prepared from flour by a dry air
classification method and by acidic water extraction. Figure 3
shows a simplified diagram of the process to be used to generate
protein concentrate from glandless cottonseed.
Three major techniques have been reported for making
glandless cottonseed isolates. These isolates can be conven-
tional protein isolates, storage proteins (SP) (soluble at high
and low pH), and nonstorage proteins (NSP) (soluble at near
neutrality):
1. Ground flour is first extracted with dilute alkali (at pH 10)
and the insoluble residue removed by continuous centrifug-
ing or decanting. The clarified liquor is then precipitated at
one pH (5.0). The resulting solids are concentrated by
centrifugation and then dried to produce a mixture of stor-
age protein and nonstorage protein.
2. In the selective precipitation procedure the protein is
precipitated with alkali at a pH 10, the solids are then
Figure 3 Protein concentrate production from glandless cottonseed. (Data from Lusas EW and Jividen GM (1987) J. Am. Oil Chem. Soc. 64(6):
839854.)
THE OILSEEDS | Cottonseed: Overview 245
separated by centrifugation. The solids in the liquor are
also precipitated at pH 7, and the storage protein curd
removed by centrifugation and then dried. The liquor is
further acidified to pH 4 to precipitate the NSP, also
removed by centrifugation and then dried. The soluble
matter remaining in the whey is then removed by either
the selective extraction or selective precipitation method.
A relatively pure storage protein fraction, containing over
90% protein (dry weight basis), can be prepared.
3. The selective extraction procedure consists basically of
leaching the proteins soluble with water at neutral pH,
where the solids are then centrifuged out. The liquor is
then acidified to pH 4; this results in a precipitated protein
curd. The curd is dried to produce an NSP isolate. The solids
from the original water leaching are then solubilized in
alkali at pH 10 and centrifuged to remove insolubles. The
liquor is then precipitated at pH 7 and the resulting curd is
concentrated and dried to produce an SP isolate.
A patented process for the preparation of glandless cottonseed
protein concentrate from solvent-extracted flour using industrial
membranes has also been used. Defatted flour is sieved through
an 80-mesh screen to break up any agglomerates and remove
hull particles, and suspended in acidified water at pH 4.04.5.
This solution is passed through an ultrafiltration membrane
(100 000 MW cutoff) and the solids retained are either dried
directly or neutralized before drying. The liquid fraction perme-
ate may then be passed through a reverse osmosis membrane to
concentrate the soluble solids and to recover water which may
be reused in the process.
Feed Uses
Glandless cottonseed has been reported to be suitable for
monogastric animal feed not just for its lack of gossypol but
also for higher available energy. The lower heat treatment
required for glandless cottonseed during milling is reported
to degrade less protein than the prepress method normally
246 THE OILSEEDS | Cottonseed: Overview
used with glanded cottonseed. Commercial glanded cotton-
seed meal intended for poultry feeding usually receives consid-
erable heat during processing to intentionally bind the
gossypol. Also in order to increase the amount of glanded
meal gossypol, addition of iron to the meal is used to inactivate
gossypol. In the cases of glandless cotton meal, it would be
expected that lower heat treatment needed for glandless cotton
would have beneficial effects in the protein availability and
quality of the meal. Some heat pretreatment of the meal is still
recommended, however, before hexane extraction, to improve
protein efficiency. Heating glandless cottonseed with 12%
added water for 10 min at 82  C, followed by 105  C for
20 min before hexane extraction, was reported to appreciably
increase broiler weight gains when compared with glanded
cottonseed feed. It has also been shown that meal from gland-
less cottonseed had an equal protein availability to soybean
meal in supporting chick growth. Feed trials with glanded
cottonseed meal with swine, catfish, fish, and shrimp have
also been described.
Exercises for Revision
What is the meaning of Glandless in relation to
cottonseed?
What is gossypol?
What natural advantage does gossypol offer to the
cotton plant?
Where is gossypol located in the cotton plant?
What is the toxicity of gossypol to humans, to animals
and to insects?
Describe the ramifications of breeding glandless cotton for
the cotton-production industry, for the cotton-oil industry
and for the food and feed industries.
Explain the relative advantages (and disadvantages) of
glandless cotton (versus glanded cotton) to all parts of the
cotton industry.
List the food uses of cottonseed and compare these with the
food uses of the soybean.
Exercises for Readers to Explore the Topic Further
Find out about the differences between the three major
species of cotton (Gossypium hirsutum L., G. aboreum L.,
G. barbadense L., G. herbaceum L.). What potential advan-
tages might accrue from the transfer of genes from one of
these to the more common cotton species?
Consider the possibility of extracting gossypol from glanded
cottonseed for use as an insecticide for growing glandless
cotton.
Research the uses and value of lecithin, and examine the
possibility of producing lecithin from glandless cottonseed.
Find out about genetically modified (GM) cotton,
the nature of its consequent tolerance to insect attack, its
suitability for producing cotton, oil and meal, and the
possible public resistance to cotton as a genetically modi-
fied organism (GMO).
See also: Agronomy of Grain Growing: Soybean: Agronomy;
Barley, Rice and Maize Processing: Oil from Rice and Maize;
Fats: Healthy Fats and Oils; Lipid Chemistry; Food Grains and the
Consumer: Genetically Modified Grains and the Consumer; Grains
Around the World: Grain Production and Consumption: Oilseeds in
North America; Non-food Products from Grains: Biodiesel; Cereal
Grains as Animal Feed; Oilseed and Legume Processing:
Soybean: Processing; Soybean: Soy Concentrates and Isolates; The
Basics: Grain and Plant Morphology of Cereals and how characters
can be used to identify varieties; Taxonomic Classification of Grain
Species; The Grain Crops: An Overview; The Oilseeds: Oilseeds:
Overview; Soybean: Overview.
Further Reading
Berot S, Chenu M, Coutret J, and Gueguen J (1995) Cottonseed protein-rich products
from glandless African varieties: 1. Pilot-plant scale production of protein
concentrates, Sciences-des-Aliments 15(3): 203215.
Camire ME, King CC, and Bittner DR (1991) Characteristics of extruded mixtures of
cornmeal and glandless cottonseed flour. Cereal Chemistry 68(4): 419424.
Cater CM (1968) Processing glandless cottonseed. Oil Mill Gazeteer 72(7): 815.
Cherry JP, Gray MS, and Jones LA (1981) A review of lecithin chemistry and glandless
cottonseed as a potential commercial source. Seed-oil content of glanded and
glandless, cottons. Journal of the American Oil Chemists Society 58(10): 903913.
Glandless cotton: Its significance, status, and prospects: Proceedings of a conference,
December 1314, 1977, Dallas, Texas/sponsored by Agricultural Research Service,
United States Department of Agriculture and National Cottonseed Products
Association, Inc., Agricultural Research Service Southern Region, US Dept. of
Agriculture, 1978.
Kohel RJ and Lee JA (1984) Genetic analysis of Egyptian glandless cotton. Crop
Science 24(6): 11191121.
Lawhon JT, Cater CM, and Mattil KF (1977) Journal of the American Oil Chemists
Society 54: 75.
La-Rue DC, Knabe DA, Izard RS, and Tanksley TD (1986) Glandless cottonseed kernels
for starter, grower and finisher swine. Journal of Animal Science 64(4): 10511059.
Lusas EW and Jividen GM (1987) Glandless cottonseed: A review of the first 25 years of
processing and utilization research. Journal of the American Oil Chemists Society
64(6): 839854.
Miravalle RJ (1972) The plant geneticists contribution toward changing the lipid and
amino acid composition of cottonseed. Journal of the American Oil Chemists
Society 49: 2426.
Percival AE, Wendel JF, and Stewart JM (1999) Taxonomy and Germ plasm resources.
In: Smith CW and Cothren JT (eds.) Cotton. New York: Wiley.
Robinson EH, Rawles SD, Oldenburg PW, and Stickney RR (1984) Effects of feeding
glandless or glanded cottonseed products and gossypol to Tilapia aurea.
Aquaculture 38(2): 145154.
Ryan JR, Kratzer FH, Grau CR, and Vohra P (1986) Glandless cottonseed meal for laying
and breeding hens and broiler chicks. Poultary Science 65(5): 949955.
Simmons RG and Golightly NH (1981) Cottonseed Cookery. College Station, TX: Food
Protein R&D Center, Texas A&M University.
Silva-Ramos LC and Kohel RJ (1987) Seed-oil content of glanded and glandless
cottons. Journal of the American Oil Chemists Society 64(9): 13371340.
Relevant Website
cottonseed.com http://www.cottonseed.com National Cottonseed Products
Association (NCPA) is a trade association for the cottonseed processing industry.
Besides commercial information, it has further links to technical information on
cottonseed products.
 Sunflower: Overview
GJ Seiler and TJ Gulya, USDA-ARS, Northern Crop Science Laboratory, Fargo, ND, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Sunflower history origin and crop development
Sunflower botany
Sunflower production practices in North America
Sunflower agronomic problems: insect, disease, and preda-
tors (birds)
Sunflower processing and uses
Learning Objectives
To understand the basics of the origin of sunflower and its
agronomy, production, processing, and uses
To understand the diversity of sunflower types and where
sunflower is produced
To understand the variation in fatty acid composition and
its significance in sunflower utilization
Overview
Sunflower (Helianthus annuus) is grown as an oilseed crop world-
wide in temperate and subtropical climates in 72 countries.
Among oilseeds, sunflower generally ranks fifth behind soybean,
rapeseed, cottonseed, and peanut, with an average annual world
production of 3244 million tonnes (Table 1). Unlike soybean,
sunflower is primarily an oil crop, with high-protein meal being
a by-product. Sunflower is grown on every continent, except
Antarctica, with Ukraine being the largest producer, followed
by the Russian Federation, Argentina, China, France, Romania,
Bulgaria, Turkey, Hungary, and the United States in 201112
(Table 2). US production accounts for 35% of the world out-
put. Nutritionally, sunflower oil is somewhat superior to other
vegetable oils due to the greater proportion of the unsaturated
fatty acids (oleic, linoleic, and linolenic) and lower saturated
fatty acids (palmitic and stearic), especially in the recently devel-
oped midoleic content NuSun hybrids (Table 3). Sunflower
oil contains zero trans fats, which have been implicated in ele-
vated cholesterol levels and increased risk of coronary heart
disease.
History
Sunflower was domesticated by Native Americans in the
Eastern United States in about 3000 BC. They used the seeds
directly as food and crudely extracted the oil. Native Americans
had selected a tall, single-headed variety by the time European
explorers reached North America in the sixteenth century.
While sunflower was not the staple that the three sisters
(maize, beans, and squash) were, it nonetheless was cultivated
by many tribes from eastern North America through the
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00027-9
Midwest and as far as northern Mexico. The Native Americans
also used sunflower hulls as a source of dye, leaves for herbal
medicines, and pollen in religious ceremonies.
Historical records indicate that the Spanish were the first to
introduce sunflower to Europe in the early 1500s. Sunflower
was initially grown as an ornamental plant. Early English and
French explorers also introduced it to their respective coun-
tries. From Western Europe, sunflower spread along the trade
routes to Egypt, Afghanistan, India, China, and Russia. By the
early 1700s, sunflower seeds were eaten as a snack, and in
1716, the first patent for the use of sunflower oil (for industrial
purposes) was filed in England. The most significant boost for
sunflower as a crop, however, came from the Russian Ortho-
dox Church. Lenten regulations prohibited the consumption
of many oily foods, but since sunflower was not specifically
listed, the seed and oil became a staple diet item in Russia.
Efforts by Russian scientists led to significant crop improve-
ments with oil contents soon exceeding 40%. While sunflower
was grown throughout Western and Eastern Europe, Russia
historically was the largest producer with in excess of 3 million
hectares in the early twentieth century, compared to one-half
million hectares for the rest of Europe. Russian immigrants are
credited with introducing sunflower to North America. In fact,
the open-pollinated variety Russian Mammoth still sold by
garden seed firms traces its lineage back to the same-named
variety initially introduced in the 1880s. Early cultivation of
sunflower in North America was primarily for livestock silage
and seed for poultry. By the second half of the twentieth
century, improved Russian varieties with oil levels of 4555%
were available. Increased US production of these high-oil sun-
flower varieties spurred interest by oil crushers, which led to
expanded US production, especially in the Northern Great
Plains (North Dakota and Minnesota). The discovery of cyto-
plasmic male sterility (CMS) by French scientists laid the foun-
dation for the development of sunflower hybrids in the early
1970s. Hybrid sunflower, with higher yields and oil content,
more uniformity, and disease resistance, in comparison with
open-pollinated varieties, provided the last great impetus in
establishing sunflower as a worldwide crop (Figure 1).
Botany
Domesticated sunflower, Helianthus annuus L., is the same
species as the wild, multiheaded common annual sunflower
found throughout the contiguous United States, northern
Mexico, and southern Canada (Figure 2). The genus Helianthus
comprises 52 species within the tribe Heliantheae of the family
Asteraceae. The only other Helianthus species grown commer-
cially is the perennial H. tuberosus, commonly known as
Jerusalem artichoke. There are 14 annual Helianthus species,
all with 2n 34 chromosomes, and 38 perennial species,
which may be diploid, tetraploid, or hexaploid. Many species
247
248 THE OILSEEDS | Sunflower: Overview

Table 1 World production (million tonnes) of major oilseed crops over the last 5 years

Commodity 2007/08 2008/09 2009/10 2010/11 2011/12

Soybean 260 264 239 268 287

Rapeseed 61 61 61 63 70
Cottonseed 39 44 48 46 44
Peanut 36 39 38 39 39
Sunflower 32 34 41 36 44
Palm kernel 12 13 14 14 15
Coconut 6 6 6 6 6
Total 446 461 447 472 505

Source: http://apps.fas.usda.gov/psdonline/circulars/oilseeds.pdf.

Table 2 World sunflower seed production (in 1000 tonnes) over the last 5 years

Country 2007/08 2008/09 2009/10 2010/11 2011/12

Ukraine 6526 6364 6771 8670 8387

Russian Federation 7350 6454 5345 9697 7993
Argentina 4650 2483 2220 3672 3340
China 1791 1955 2298 2313 2369
France 1598 1716 1635 1880 1573
Romania 1170 1098 1263 1789 1398
Bulgaria 1300 1317 1536 1439 1387
Turkey 992 1057 1320 1335 1370
Hungary 1468 1265 970 1374 1316
The United States 1552 1377 1240 924 1263
Others 7852 7793 6891 7620 7053
Total 36 249 32 878 31 489 40 713 37 449

Source: http://faostat.fao.org/.

Table 3 Fatty acid composition of traditional, high oleic and NuSun sunflower oil in comparison with other vegetable oils

Saturated fatty acids Unsaturated fatty acids

Crop
Palmitic and stearic Oleic Linoleic Linolenic

Sunflower, traditional 12.5 20 69 0.1

Sunflower, high oleic 9 82 9 0.1
Sunflower, NuSun 9 65 26 0.1
Olive 17 72 11 1
Rapeseed 6 62 22 10
Soybean 15 24 61 7
Corn 13 25 61 1

Source: http://www.sunflowernsa.com.

freely hybridize with each other in nature, giving rise to inter-
mediate forms, which has led to confusion in taxonomy. While
H. annuus is a geographically diverse species occupying a wide
range of habitats, other Helianthus species are very habitat-
specific and thus are endemic in a limited number of locations.
For example, H. niveus ssp. tephrodes is only found in active
sand dunes in southern California and in northern Sonora,
Mexico. H. paradoxus is only found in saline, marshy sites in
western Texas and New Mexico, and H. exilis is only found in
serpentine sites in Northern California whose soils are nearly
toxic to many nonadapted plants. Wild Helianthus species,
both annual and perennial, have contributed many useful
traits to domesticated sunflower, including disease resistance,
drought and salt tolerance, herbicide resistance, CMS, and
improved oil quality. All annual species, except H. agrestis,
can be crossed with cultivated H. annuus. There has been
more success in crossing tetraploid and hexaploid perennial
species with cultivated H. annuus than with the diploid peren-
nial species.
Production Practices in North America
Sunflower production in the United States is concentrated in
the Northern Great Plains (North Dakota, South Dakota, and
Minnesota) with a secondary production area in the Central
Great Plains (western Kansas, Nebraska, and eastern Colorado)
(Table 4), but there is some amount of sunflower grown in
 THE OILSEEDS | Sunflower: Overview 249

Table 4 Sunflower production (hectares planted) and yield by state

in the United States over the last 3 years

Production ( 1000 ha) Yield (kg ha1)

US state 2010 2011 2012 2010 2011 2012

North Dakota 272 202 305 1702 1530 1663

South Dakota 162 163 227 2016 1863 1451
Kansas 43 43 26 1769 1372 1372
Minnesota 21 11 15 1568 1395 1941
Colorado 37 39 25 1478 1231 876
Nebraska 10 14 12 1343 1574 1022
Texas 11 9 13 1007 998 1242
Other 15 18 21 1343 1140 1341
Total 571 499 644 1750 1566 1516

Figure 1 Sunflower field in bloom. Courtesy of Dr. Brady Vick (Retired),
USDA-ARS Sunflower and Plant Biology Unit, Fargo, ND, USA.
Figure 2 Common annual sunflower, Helianthus annuus, the
progenitor of cultivated sunflower, with multiple flowers on a branched
stem. Courtesy of Dr. Brady Vick (retired), USDA-ARS Sunflower and
Plant Biology Unit, Fargo, ND, USA.
each of the 48 contiguous states (especially for birdseed). Most
of the production recommendations have come from research
conducted in North Dakota, but as interest in sunflower
increases in other areas, new research generated from other
states addresses these different environments.
Sunflower is traditionally planted between May 1 and the
middle of June in the Northern Great Plains, with hybrid
maturities of 100120 days to fit the short, 120-day average
frost-free growing season of this region. Longer maturing
hybrids are used where short growing seasons are not a restric-
tion. Similarly, there are early maturing hybrids that would be
suitable for a double-crop situation. Double-crop sunflowers
are typically planted in July, either following the failure of an
initial crop or after harvest of a small grain crop. Yields of
22003500 kg ha1 are attainable under optimal, irrigated
conditions, while the average dryland yield of sunflower in
the United States ranges from 1516 to 1750 kg ha1. The phe-
nology of sunflower hybrid maturation is depicted in Table 5.
Source: http://www.sunflowernsa.com/.
Sunflower is typically planted when soil temperatures reach
10  C. Below 10  C, germination is very slow. Sunflower is gen-
erally planted in row widths of 5075 cm to match the available
equipment, although solid seeding (widths < 30 cm) is also
practiced. Sunflower is planted from 4 to 7 cm deep at rates of
3762 000 seeds ha1 for oilseed and 3550 000 seeds ha1 for
confection. Lower plant populations for confectionery sunflower
produce larger-sized heads and a greater proportion of large seed,
which is desirable. Conversely, high plant populations will result
in smaller heads, which not only dry down faster but also have
thinner stalks that are more prone to lodging.
Sunflower has a root system characterized by both a deep
taproot and an extensive fibrous root system, which makes it
efficient at both water uptake and nutrient uptake. While its
water requirements are similar to that of corn, sunflowers greater
efficiency can result in higher returns under moisture-limiting
conditions. Sunflower responds well to nitrogen fertilization,
with a general rule of 5 kg ha1 nitrogen required for 100 kg
seed ha1. Thus, for a yield target of 2000 kg seed ha1, the soil
nitrogen content plus added nitrogen would be 100 kg ha1.
Early weed control is important in sunflower with several
preplant, preemergence herbicides available for grass control
and now postemergent ones for broadleaf weed control.
Hybrids have been developed through conventional breeding
with tolerance to both imidazolinone and sulfonylurea herbi-
cides, allowing for postemergent applications. There are no
glyphosate-resistant hybrids in any country, nor are there
currently any other commercially available traits introduced
into sunflower from other organisms beyond the genus
Helianthus (i.e., sunflower is currently still a non-GMO crop).
Sunflower in the Northern Great Plains typically flowers in
August, about 6070 days after planting. Four to five weeks
post-pollination is required for seed maturation, at which
point seed moisture is 3040%. If the crop is killed by frost,
seeds may be harvested without use of a chemical desiccant.
Any conventional grain combine can be used to harvest sun-
flower, with a variety of attachments to facilitate harvesting of
the sunflower heads. Chemical desiccants can be used to
reduce seed moisture, minimizing losses due to birds, lodging,
and severe disease attack of the heads. Seed moisture must be at
10% or less to retard fungal deterioration in storage, with 8%
moisture preferable for long-term storage.
250 THE OILSEEDS | Sunflower: Overview

Table 5 Sunflower growth stages and approximate time and growing degree days (GDD) to reach each stage

Sunflower growth stage Description GDD Days to reach stage

VE Emergence 93 10
V4 4 True leaves 194 20
V8 8 True leaves 303 28
V12 12 True leaves 383 34
V16 16 True leaves 429 38
V20 20 True leaves 484 44
R1 Miniature terminal bud 511 46
R2 Bud <2 cm from last leaf 695 61
R3 Bud >2 cm from last leaf 774 67
R4 Bud open, ray flowers visible 829 71
R5.1 Early flower 859 73
R5.5 50% flowered 902 77
R6 Flowering complete 989 84
R7 Back of head pale yellow 1140 86
R8 Bracts green, head back yellow 1228 104
R9 Bracts yellow, head back brown 1372 119

(GDD ((Max. temp. in  C)  (Min. temp. in  C)/2)  10  C).

Source: NDSU Extension Bulletin 25 (revised). Sunflower Production.

Pest Problems
Cultivated sunflower has a number of pest problems, including
diseases, insects, and birds. Since the genus Helianthus is indig-
enous to North America and coevolved with the pests, there are
a great diversity of insects and pathogens adapted to sunflower,
plus a natural reservoir of these pests on the wild progenitors.
On other continents, the diversity of sunflower pests and path-
ogens is generally less, although international seed movement
has dispersed most sunflower pathogens around the globe.
Diseases
Sunflower is subject to a number of fungal, bacterial, and viral
diseases, with fungi the most numerous and economically
serious. Diseases causing the most losses worldwide are Scler-
otinia head rot and stalk rot, Phomopsis stem canker, rust, and
downy mildew. Some diseases are serious in only a few coun-
tries, such as Verticillium wilt in Argentina or white rust
(Albugo) in South Africa. Most sunflower diseases are caused
by pathogens specific to sunflower, such as Phomopsis helianthi,
Alternaria helianthi, and Plasmopara halstedii. Some of the most
serious diseases, however, are caused by pathogens with wide
host ranges, such as Sclerotinia sclerotiorum that occurs on 137
different genera of plants.
Disease organisms may be grouped based on plant parts
affected. The only significant seedling disease is downy mildew
(Plasmopara halstedii). Foliar diseases on sunflower include rust
(Puccinia helianthi), blights caused by Alternaria (five species) and
Septoria helianthi, powdery mildew (Erysiphe cichoracearum), bac-
terial blight (Pseudomonas syringae pv. helianthi), apical chlorosis
(Pseudomonas syringae pv. tagetis), sunflower mosaic virus, and
sunflower chlorotic mottle virus. The main stem diseases of
sunflower are Sclerotinia stalk rot (S. sclerotiorum), stalk rots
caused by S. minor and Sclerotium rolfsii, Phomopsis stem canker
(P. helianthi), charcoal rot (Macrophomina phaseolina), and bac-
terial stalk rot (Erwinia carotovora). Head rots are probably the
most devastating because of their direct impact upon seed yield.
The main head rot pathogens are Sclerotinia sclerotiorum, Rhizopus
(three species), Botrytis cinerea, and Erwinia carotovora.
Control of most diseases is accomplished primarily with
resistant hybrids and, to a lesser extent, with cultural practices.
Single dominant genes are used to confer resistance to rust,
Verticillium wilt, and downy mildew. At least two dominant
genes are required for resistance to Phomopsis stem canker,
while resistance to Sclerotinia head rot and stalk rot is con-
trolled polygenically, making it the most difficult disease to
select for resistance. Very few diseases are managed with fun-
gicides, primarily because of economics. Seed treatments for
control of downy mildew had been very effective until the
pathogen developed resistance to the fungicide metalaxyl. In
some countries, foliar fungicides are used to manage Phomopsis
disease and rust. The use of foliar fungicides is more practical
with confection sunflower where the profit margin is greater
than with oilseeds.
Insects
In the United States, there is a wide variety of sunflower-
specific, native insects that preferentially feed on stems, leaves,
roots, or seeds. Stem-feeding insects on sunflower include at
least three different cutworms (Euxoa species and Feltia jaculi-
fera), the sunflower stem weevil (Cylindrocopturus adspersus), the
black sunflower stem weevil (Apion occidentale), the sunflower
maggot (Strauzia longipennis), and the long-horned sunflower
stem girdler (Dectes texanus). Root-feeding insects include the
carrot beetle (Ligyrus gibbosus), the sunflower root weevil (Baris
strenua), and occasionally wireworms. Leaf-feeding insects
include grasshoppers (four Melanoplus species and Camnula
pellucida), aphids (primarily Aphis helianthi and Masonaphis
masoni), the painted lady caterpillar (Vanessa cardui), the sun-
flower beetle (Zygogramma exclamationis), and the palestriped
flea beetle (Systena blanda). Seed- and head-feeding insects
include the sunflower moth (Homoeosoma electellum), the sun-
flower midge (Contarinia schulzi), the red sunflower seed weevil
(Smicronyx fulvus), the gray seed weevil (Smicronyx sordidus), the

banded sunflower moth (Cochylis hospes), the sunflower head
moth (Gymnocarena diffusa), the sunflower seed maggot (Neote-
phritis finalis), the sunflower headclipping weevil (Haplorhynch-
ites aeneus), and the tarnished plant bug (Lygus lineolaris). In
other continents, insects are generally considered minimal or
infrequent problems, and when they occur, they are generally
caused by omnivorous insects such as aphids, plant bugs (Lygus
spp.), and other non-sunflower-specific insects. Management
of sunflower insect pests usually concentrates on cultural and
chemical strategies. To date, selection for effective insect resis-
tance has been limited in cultivated sunflower.
Birds
Sunflower is vulnerable to damage by birds due to its easy
accessibility and high nutritional value of the seed. Ranging
from tiny sparrows (Passeridae) to large parrots (Psittacidae),
birds are a constant problem to sunflower on all continents. In
commercial birdseed mixtures, sunflower is the preferred seed,
so it is not surprising that the crop is beset by depredation by a
wide variety of birds. In the United States, the migratory red-
winged blackbird (Agelaius phoeniceus) causes the most dam-
age. In Europe, many different sparrows (Passer species) and
doves (Streptopelia spp.) are the major problems, while in
South America, parakeets (Psittacidae) and doves (Columbidae)
predominate.
Seed losses due to bird feeding can easily exceed 10% in
fields planted close to nesting areas, and losses of 100% are not
unheard of. In the United States, yield loss due to birds is
significant enough that many different management strategies
have been investigated. There has been limited success in
breeding for sunflower varieties that have physical barriers
(long necks, tight bracts, and seed coloration) to bird feeding.
Most research efforts have explored cultural methods, habitat
management, and mechanical frightening devices (noise
makers). Various registered avicides have been tested, but
none have been successful in reducing bird depredation. How-
ever, one novel approach under investigation is a commercial
product, which contains the FDA-approved grape flavor com-
ponent, which ironically many birds find repellent.
Sunflower Oil, Processing, and Uses
Sunflower is grown worldwide, primarily as an edible oil crop.
Traditional sunflower oil (high linoleic) is used mainly as salad
oil and frying oil, although it can be hydrogenated for use in
margarines. In terms of world production, sunflower usually
ranks fifth among oilseed crops behind soybean, rapeseed or
canola, cottonseed, and peanut. The oil content of oilseed
sunflower varies from 40% to 50% by weight. There are cur-
rently three categories of sunflower oil: traditional, NuSun,
and high oleic. Traditional sunflower oil is characterized by a
high concentration of linoleic acid (70%) and a moderate
amount of oleic acid (20%). Plant breeders, using conven-
tional methods, have selected lines in which oleic acid is the
major fatty acid component (65%), and this oil has been
given the trade name of NuSun. High oleic oil has an oleic
acid content of 8590%. Oleic acid, a monounsaturated fatty
acid, is considered by health and dietary experts to be better
THE OILSEEDS | Sunflower: Overview 251
nutritionally than polyunsaturated fatty acids, like linoleic
acid, and far superior to saturated fatty acids. While oleic acid
levels of 8090% are attainable, the frying industry prefers an
oleic acid content of 5560%, primarily because of the taste
associated with the remnant linoleic acid. High and mid-oleic
sunflower oil also has a very high oxidative stability, almost
10  that of traditional sunflower oil, extending product shelf
life and longevity of the frying oil. For a complete comparison
of the fatty acid composition of various vegetable oils (see
Table 3).
Oil extraction from sunflower seed is a relatively straight-
forward process. Seed is first thoroughly dried, dehulled, and
then flaked or rolled and heated to 8590  C. The crushed seed
is then subjected to one of three extraction methods: (1) a
mechanical screw press, (2) extraction with an organic solvent,
usually hexane, followed by distillation to remove the solvent,
and (3) a combination of the screw press and solvent extrac-
tion. In the United States, the latter is the preferred method and
generally recovers about 99% of the total seed oil. The crude oil
is further processed to remove gums, waxes, free fatty acids,
and odors. The refined oil is now suitable as a vegetable oil or
can be hydrogenated and blended with soybean or canola oil
for margarines.
Sunflower oil can be used as an alternative or additive to
diesel fuel to create biodiesel, a clean burning alternative fuel
produced from a renewable resource. This could help farmers
decrease their dependence upon petroleum fuels by substituting
farm-grown fuel. For diesel engine use, sunflower oil requires
more extensive purification including removal of waxes and
gums. Minor engine modifications, including improved fuel
filters, are also necessary to burn any vegetable oil. Since the
energy content of sunflower oil is less than that of diesel, fuel
consumption would be greater and power output less. As the
price of diesel increases and the world supply diminishes, the
feasibility of vegetable oil use in diesel engines becomes more of
a reality.
Sunflower By-products
Sunflower Meal
After sunflower seeds are crushed, the remaining meal is a high-
protein product commonly used in livestock and poultry feeds.
The meal contains 260500 g kg1 protein, 120350 g kg1
fiber (from remnant hulls), and 1090 g kg1 fat. Sunflower
meal is lower in lysine content and higher in methionine than
soybean meal and thus is usually combined with soybean meal
in livestock feeds. Feeding trials with both dairy and beef cattle
have adequately demonstrated the utility of sunflower meal
since the meal is more ruminally degradable than either soybean
or canola meal.
Hulls
Sunflower hulls comprise 2130% of the total seed weight and
are often a waste by-product. In oil extraction factories, the
hulls are often burned as a source of heat, both for the plant
and for the distillation process to remove the hexane solvent.
Hulls, mixed with appropriate binders and glue, have also been
molded into cylinders and sold as ersatz fireplace logs. Public
252 THE OILSEEDS | Sunflower: Overview
buildings in the vicinity of processing plants have also used
sunflower hulls for heating, with only minor modifications of
their boilers.
A red dye, suitable as an FDA-approved food colorant, can
be extracted from the hulls of purple-seeded sunflower. With
the trend away from the use of synthetic dyes, many foods high
in anthocyanin content have been viewed as dye sources. The
anthocyanin content of purple-hulled seed ranges from 6 to
16 g kg1 of hulls, which compares favorably with that of other
sources (grape skins and beet pulp). Natural red dyes lack the
chemical stability of synthetic dyes and thus may only be
usable in certain applications.
Pectin, used in jellies and as a general food thickener, can be
extracted from sunflower heads. The pectin content, after seed
removal, is from 150 to 250 g kg1, and thus, a hectare of
sunflower could yield 200350 kg pectin. Sunflower pectin is
chemically a low-methoxyl pectin, as contrasted to high-
methoxyl pectins in apples and citrus. Low-methoxyl pectins
are used in making jellies low in sugar and thus would be used
primarily in the diet food market.
Sunflower butter, a relatively new product marketed as
SunButter, is a delicious and healthy alternative to peanut
butter. Packed with nutrition, it is an excellent choice for
people with peanut allergies. Made from specially roasted sun-
flower seeds, it is completely peanut-free, tree nut-free, and
gluten-free. A single serving of SunButter Natural Omega-3
provides 461 mg of omega-3, with 1/3 less saturated fat than
peanut butter.
Confection Sunflower
Sunflower hybrids grown for human consumption are termed
confectionery sunflower. The seeds are generally roasted,
salted, and eaten as a snack food, either in-shell or dehulled
to produce nutmeats. The nutmeats are also used in breakfast
cereals, trail mixes, and baked goods. Sunflower kernels have
been chocolate- or candy-coated as a snack. In shell, they have
also been coated with various flavorings, such as bacon, bar-
beque, salt, and pepper to name a few. Confectionery
sunflower comprises 1822% of US total sunflower produc-
tion, with the United States the largest producer in the world
market, followed by Argentina. Confectionery sunflower is
characterized by its larger seed size and by having white or
gray stripes on a black or brown background, as compared to
uniform black color of oilseed sunflower.
Confectionery seed is graded on the basis of size. The largest
seeds, referred to as in-shell, are those passing over a 7.9 mm
round-hole screen and generally make up 1525% of the
harvested seed. The medium-sized seed, destined for dehulling
for use as nutmeats, comprises 3050% of the crop and is that
fraction recovered between a 7.9 and 7.1 mm round-hole
screen. The smallest seeds, passing through the 7.1 mm
round-hole screen, are used primarily as birdseed and com-
prise 1520% of the crop.
Confectionery sunflower is genetically unique from oilseed
sunflower, primarily because seed size and hull coloration are
more important than oil content. As a group, they are much more
susceptible to diseases and insects. Since the profit to the grower
may be 25100% greater than with oilseeds, extra management
practices to control insects and diseases are more feasible.
Future Trends
Sunflower production in the United States is in direct compe-
tition with soybean and rapeseed, which currently have the
advantage of glyphosate resistance for weed management. Sun-
flower is also confronted by disease and insect problems that
are lesser concerns in soybeans and rapeseed. The challenge to
the sunflower industry is to make the crop as easy and profit-
able to grow as its competitors. The development of
imidazolinone- and sulfonylurea-resistant hybrids gives
growers new tools to control postemergent weeds, similar to
the glyphosate resistance technology. Finding management
schemes to control insects and diseases in sunflower is a con-
tinuing challenge. Worldwide, Sclerotinia and Phomopsis
diseases are the two most devastating diseases. Both diseases
can be managed to some degree with fungicides, but genetic
resistance would be more economical and safer for the envi-
ronment. The many native Helianthus species offer untapped
resources for resistance genes for Sclerotinia and other diseases.
Another option to make sunflower more competitive is to
change its fatty acid composition to make the oil nutritionally
better than other oils, the so-called designer-specialty oils.
Moves in this direction include the NuSun hybrids, with
oil nearly comparable to olive oil. The global sunflower market
is trending toward high oleic (8590%) oil, which has zero
trans fats, low saturated fats, and high monounsaturated fatty
acid with high oxidative stability. The transformation is already
under way but will probably take 10 years to totally convert the
industry. As the world public becomes more health-conscious
and with the trend to increase label information, consumers
are demanding to know what in a product makes it a healthy
choice. Healthy sunflower oil will benefit from these demands
and in turn generate more interest among growers. Commit-
ments by large snack food processors and fast-food chain
restaurants in the use of NuSun have been the first step in
increasing domestic usage of sunflower oil.
Exercises for Revision
From which continent did the sunflower originate?
Who introduced the sunflower to Europe?
Which are the main sunflower-producing countries?
Who were the first to increase sunflower oil content?
How many days are needed for sunflower to reach maturity
in North America?
What are the various uses of sunflower?
What are NuSun sunflowers?
Which birds are the main problems in North and South
Americas?
Exercises for Readers to Explore the Topic Further
Propose novel technological or plant breeding solutions to
the problem of bird damage in sunflower production.

Compare the productivity of sunflower with that of other
oilseed crops in a region of interest to you.
What scope is there for further value-added development of
specialty oils from sunflower?
Further Reading
Berglund DR (2007) Sunflower Production. Fargo, ND: North Dakota State University
Extension Service Bulletin 25 (revised), 119 pp.
Fick GN (1989) Sunflower. In: Robbelen G, Downey RK, and Shri A (eds.) Oil Crops of
the World. New York: McGraw-Hill, 553 pp.
Heiser Jr. CB (1976) The Sunflower. Norman, OK: University Oklahoma Press, 198 pp.
Heiser Jr. CB, Smith DM, Clevenger SB, and Martin Jr. WC (1969) The North American
sunflowers (Helianthus). Memoirs of the Torrey Botanical Club 22(3): 1218.
Jan CC and Seiler GJ (2007) Sunflower. In: Singh RJ (ed.) Genetics Resources,
Chromosome Engineering, and Crop Improvement. Oilseed Crops, vol. 4.
New York: CRC Press, Taylor and Francis Group Chapter 5, 304 pp.
Maiti RK, Singh VP, Purohit SS, and Vidyasagar P (eds.) (2007) Research Advances in
Sunflower. Jodhpur, India: Agrobios International, 512 pp.
Miller JF (1987) Sunflower. In: Fehr WR (ed.) Principles of Cultivar Development, vol.
2. New York: Macmillan Publication, 761 pp.
Pappalardo J (2008) Sunflower, the Secret History. Woodstock, NY: The Overlook
Press, 256 pp.
Rogers CE, Thompson TE, and Seiler GJ (1982) Sunflower Species of the United States.
Bismarck, ND: National Sunflower Association, 75 pp.
THE OILSEEDS | Sunflower: Overview 253
Salunkhe DK, Chavan JK, Adsule RN, and Kadam SS (1992) Sunflower. In: World
Oilseeds: Chemistry, Technology, and Utilization. New York: Van Nostrand
Reinhold, 554 pp.
Schneiter AA (1997) Sunflower Technology and Production. Madison WI: American
Society of Agronomy Monograph #35.
Seiler GJ (ed.) (1992) Sunflower: Special Issue. Field Crops Research 30(34):
191448.
Seiler GJ and Jan CC (2010) Basic sunflower information. In: Hu J, Seiler G, and Kole C
(eds.) Genetics, Genomics, and Breeding of Sunflower, pp. 150. Enfield, NH:
Science Publishers Chapter 1, 335 pp.
Skoric D, Seiler GJ, Zhao L, Jan CC, Miller JF, and Charlet L (eds.) (2012) Sunflower
Genetics and Breeding. Novi Sad, Serbia: Serbian Academy of Science and Arts,
520 pp.
Weiss EA (ed.) (2000) Sunflower. In: Oilseed Crops, 364 pp. Malden, MA: Blackwell
Science.
Relevant Websites
www.ars.usda.gov/main/site_main.htm?modecode54-42-05-20 U.S. Department
of Agriculture Agricultural Research Service, Sunflower and Plant Biology
Research Unit.
http://www.ext.nodak.edu/extpubs/plantsci/rowcrops/eb25w-12.htm North Dakota
State University Extension Service, with individual bulletins on many aspects of
sunflower production.
www.isasunflower.org International Sunflower Association.
www.sunflowernsa.com National Sunflower Association.
 Overview of the Oilseed Safflower (Carthamus tinctorius L.)
C Hall III, North Dakota State University, Fargo, ND, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Carthamus is part of the Compositae or Asteraceae family.
Carthamus tinctorius L. is the cultivated species of safflower.
Flowers of safflower are used as a food colorant and cloth-
ing dye and in traditional Chinese medicine.
The oil from safflower is 90% oleic and linoleic acids and is
relatively stable to oxidation.
Safflower contains high levels of flavonoids and
tocopherols.
Oil composition can be manipulated to create health-
promoting oils or biofuel feedstock.
Safflower is an oilseed used in commercial bird foods.
Defatted safflower meal can be used as a cattle feed.
Learning Objective
To gain knowledge in the fundamental aspects of safflower
production, composition, and uses.
Introduction
The Food and Agriculture Organization considers safflower
(Carthamus tinctorius L.) as a minor oilseed crop since only
about 647 000 MT, on 783 000 Ha, is produced annually. Saf-
flower can be produced on arid and semiarid land, including
moderate salinity, and is therefore produced across the world
where water is scarce or high salinity water is used for irriga-
tion. Carthamus tinctorius L. is the cultivated species of saf-
flower. The word tinctorius essentially means for dyeing in
English. The flowers of safflower have been used historically
as a colorant in food and as dye in the clothing industry and as
a traditional Chinese medicine. However, the primary purpose
today is as an oilseed crop. The oil is typically used for cooking
and frying, but interest as a feedstock for biodiesel is growing.
Safflower oil composition varies widely, but 90% of the oil is
made up of oleic and linoleic acids. However, the genes of
Carthamus species can be easily manipulated to produce oils
with unique fatty acids, which can then be used for specific
purposes. In North America, much of the safflower is used as
wild bird feed. This article provides the most recent informa-
tion on safflower, and readers are encouraged to review mate-
rials found in the referenced materials, as sources for historical
and early agronomic information.
Taxonomy, Morphology, and Agronomics
A number of locations have been proposed as the center of
origin for safflower. These include Central Asia and the Middle
East. Weiss provided an excellent historical perspective of
254 Encyclopedia of Food Grains, Second Edition
safflower, which dates to 2000 BC in Egypt. The height of
safflower production occurred shortly after World War II and
has since been replaced with other oilseed crops. Safflower
production ranks eighth among oilseeds.
The evolutionary history of safflower has been documen-
ted to approximately 1753. Although C. tinctorius L. is the
only cultivated species, there were as many as 25 species
linked to the genus Carthamus. However, clarification over
the last two decades supports the presence of approximately
15 species in the genus Carthamus. Much of the confusion
around this genus was due to limited morphological obser-
vation such as spiny leaves or cypselas. The advent of molec-
ular biology allowed for the differentiation of Carthamus
species.
Carthamus species are grouped into sections based on chro-
mosome pairs. Species with 12 chromosomal pairs include
C. tinctorius, C. palaestinus, and C. oxyacantha, while other species
contain 10, 11, 22, and 32 chromosomal pairs. C. tinctorius,
C. persicus Desf. Ex Willd, C. palaestinus, and C. oxyacantha have
12 chromosomal pairs and can readily cross-pollinate. Only
C. tinctorius is commercially produced, while the other three
species are wild. However, these species can cross-pollinate
resulting in altered seed characteristics. C. tinctorius also can
cross with wild species having 10 and 22 chromosomal pairs
but not 32. The crossing of C. tinctorius with weedy Carthamus
species (e.g., C. lanatus) has been documented. However, the
propensity for weediness in the resulting F1 hybrid was less than
the parent wild species and no F2 hybrid (i.e., result of crossing
of two F1 hybrids) was produced.
A number of factors contribute to the growth characteristics
of Carthamus species. However, much of the plant breeding
efforts have focused on C. tinctorius L. Therefore, only this
commercially relevant species will be presented in the remain-
ing part of this article. Safflower is a thistle-like (Figure 1) plant
with yellow, orange, red, or white flowers. Leaves of the mature
plants are spiny in nature and thus compared to a thistle, while
the seeds resemble a small sunflower (Figure 2). Germination
of seeds occurs readily (i.e., 34 days) above 60  F, but growth
slows during the rosette (i.e., a circular pattern of growth) stage
that lasts up to 3 weeks. The development of a tap root, which
can reach lengths of 305 cm, occurs during the rosette stage.
This stage of growth is followed by a profuse branching stage of
growth. Overall growth behavior is dictated by the plant pop-
ulation, where bush-type plants result from greater spacing,
while closer spacing results in an upright plant form. Typical
C. tinctorius L. plant heights range from 42 to 113 cm. How-
ever, plants can reach heights of 251 cm in Asia. The plant
growth behavior also impacts seed yield. Bush-type plants
yield greater numbers of flowers as each branch typically has
five flowers with up to 50 flower heads per plant. Harvesting of
seeds usually begins 30 days after maturing and seed moisture
should not exceed 8% moisture. Much of the current breeding
focus has been directed at improving oil composition.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00030-9
 THE OILSEEDS | Overview of the Oilseed Safflower (Carthamus tinctorius L.)
Figure 1 Safflower (Carthamus tinctorius L.) in bloom. Photo
courtesy of Dr. Hans Kandel, Extension Agronomist Broadleaf Crops,
North Dakota State University.
Figure 2 Safflower (Carthamus tinctorius L.) seed.
Both cultivated safflower and wild species grow in temper-
ate zones under arid and semiarid conditions. Safflower can be
grown under diverse conditions, such as drought and salinity,
due to the long taproot accessing deep soil moisture. Contin-
ued research in area further supports the resilience of this
plant. Although safflower can grow under water-restricted envi-
ronments, Kizil et al. reported that higher amounts of rain
significantly increased a number of agronomic characteristics
of safflower grown over a 2-year period.
Composition
Protein, Fiber, Oil, and Fatty Acids
Safflower is composed of approximately 3345% hull and the
remaining fraction being the kernel. Oil content is variable,
ranging from 27% to 60% in thick- and thin-hulled cultivars,
respectively. The oil is located primarily in the kernel, where
255
greater than 95% of the oil is found. The kernel typically
contains approximately 60% oil and 26% protein. However,
the protein content of whole seed ranges from 14% to 23%.
Fiber content ranges from approximately 11% in thin-hulled
cultivars to as high as 34% in thick-walled cultivars.
Most commercial safflower oil consists of 7278% linoleic
acid and represents cultivar and environmental effects on oil
composition. Oleic acid is the second most common fatty acid
and accounts for 1520% of the fatty acids in safflower oil.
Other fatty acids include palmitic and stearic at levels ranging
from 1.5% to 6%. Only a small (<0.2%) amount of linolenic
acid exists in safflower oil. Cultivar (i.e., genotype) is one of the
drivers for the diverse oil content and oil fatty acid composi-
tion of safflower. Environment also contributes to alteration in
oil content and composition. In recent years, modification of
genes through molecular approaches has also been used to
change oil and fatty acid contents.
Crossing of different C. tinctorius genotypes produces plants
with varying oil and fatty acid compositions. Oil contents of
2533% for eight experimental safflower genotypes have been
reported. Linoleic acid represents between 56% and 75% of the
fatty acids in the oil. The genotype with the lowest linoleic acid
content had both the highest oil content and oleic acid content
(i.e., 35% of oil). Attempts to improve oil yields by crossing
C. tinctorius genotypes with wild species have had little success,
one reason being that the wild species have lower oil contents,
that is, 32%, 25%, and 17% for C. tinctorius, C. oxyacantha, and
C. lanatus, respectively, compared to greater than 40% for
cultivated safflower. However, the two wild species did have
slightly higher oleic acid contents compared to C. tinctorius.
Recently, C. tinctorius L. cv. Centennial genotype was genet-
ically altered using a cloned delta-6-desaturase gene from
Saprolegnia diclina inserted into the safflower genome by Agro-
bacterium tumefaciens. This genome manipulation resulted in a
safflower with approximately 67% gamma-linolenic acid
(GLA), which is not normally present in safflower oil. The
health benefit of GLA has been documented and is the primary
reason for the development of high-GLA safflower. In contrast,
by happenstance, a high-oleic acid safflower was identified in
1957 from seeds grown in India. The seed composition was
7781% oleic acid and 1015% linoleic, the opposite of tradi-
tional safflower. High-oleic acid safflower is available today
due to commercial production of this cultivar.
Although a few reports indicate that temperature can
impact linoleic acid content in safflower oil, temperature dur-
ing growing appears to have less influence on linoleic acid
content of safflower compared to other oilseeds. Oleoyl phos-
phatidylcholine D12-desaturase is an enzyme that regulates
linoleic acid conversion from oleic acid. Esteban et al. reported
that this enzyme was more thermally stable and less oxygen-
dependent in safflower compared to the same enzyme in sun-
flower. The high-(i.e., 7278%)linoleic acid content observed
in safflower grown under high temperatures is contrary to
other oilseeds.
Other agronomic practices have significant bearing on agro-
nomic traits. Linoleic acid content was higher in mature seed
from the parent seed having later planting dates, while all other
agronomic traits such as oil content decreased. Several authors
observed that early planting of safflower leads to higher oil
256 THE OILSEEDS | Overview of the Oilseed Safflower (Carthamus tinctorius L.)
content. Seed planting density is another agronomic practice
used to enhance yield of many crops. However, safflower tends
to branch when sown in wide seed spacing (i.e., less seed
density) and thus compensates for the lower plant numbers
by producing more seeds per plant compared to greater seed
densities. As a result, oil yield per hectare and oil content per
seed are not significantly different among plants grown from
low- and high-density seeding rates.
Harvesting of safflower usually begins 3540 days after
flowering. These harvesting dates also coincide with maximum
oil content. Thus, harvesting date can influence oil yield. The
maturity of seed also contributes to the final composition of
the seeds. Over the course of 100 days, palmitic and oleic acids
increased, while stearic acid decreased. In contrast, linoleic acid
reached a maximum at day 75 and then decreased when mea-
sured at day 100. Safflower can grow on marginal cropland,
including soils of moderate salinity. Under hydroponic condi-
tions, 50 mM sodium chloride was found to inhibit safflower
growth and increase the oxygenated essential oils. In general,
7% and 29% reductions in oil content and oil yield, respec-
tively, in safflower subjected to saline stress have been
reported. Also, oleic acid tended to increase under saline
conditions, while linoleic decreased.
Nonsaponifiable Components
Tocopherols, carotenoids, and phytosterols make up the non-
saponifiable lipid components of safflower oil. Traditional and
high-oleic acid safflower oils are composed of approximately
250 mg tocopherol per gram of oil or 50100 mg tocopherol per
gram of seed. However, tocopherol contents as high as 684 and
941 mg g 1 of oil have been reported for traditional safflower
and high-oleic acid safflower oil, respectively. Mean tocoph-
erol concentrations from seven geographic regions in the range
of 676827 mg g 1 of oil have been reported. No differentia-
tion of safflower oil type was provided, but the lowest and
highest tocopherol contents were from East Africa and South-
west Asia geographic locations, respectively.
In traditional safflower, a-tocopherol accounts for 9398%
of the total tocopherols, while b-, g-, and d-tocopherols
account for the remaining in varying concentrations.
a-Tocopherol accounted for 8398% of the tocopherols in
high-oleic acid safflower oil; however, no b analog was
found. In addition to tocopherols, 5 and 12 mg of a- and
g-tocotrienols per gram oil, respectively, were reported in cul-
tivated safflower. In contrast to cultivated species, g-tocopherol
contents as high as 36% were reported in oil obtained from
wild safflower. A mutant safflower obtained through breeding
contained a unique tocopherol profile where g-tocopherol
accounted for 96% of the total tocopherols.
Approximately 29% of the tocopherols are removed during
refining, with the greatest removal occurring during the bleach-
ing and deodorization steps of the process. Thus, adjustments
to the aforementioned numbers should be considered when
refining safflower oil. Roasting of the safflower at temperatures
between 140 and 180  C did not negatively impact tocoph-
erols or tocotrienols.
The safflower carotenoids are present in trace amounts in
the oil. Total carotenoid levels of less than 2.5 mg g 1 oil are
common. Lutein and zeaxanthin levels are approximately 0.8
and 1.5 mg g 1 oil, respectively, as determined by HPLC. How-
ever, photometric (445 nm) analysis indicated levels as high as
7 mg g 1 seed. The underestimations of carotenoids could
occur for oils that contain predominantly carotenes. In the
case of safflower, trans- and cis-b-carotene levels were only
0.140.16 mg ml 1 oil, while a-carotene accounted for less
than 0.04 mg ml 1 oil. Therefore, the total carotenoid level
close to 2.5 mg g 1 oil is realistic. Contrary to expectations,
the yellow pigments in the flowers of safflower are not carot-
enoids. Although carotenoid levels as high as 650 mg g 1 dried
flower petals have been reported, hydroxyl safflower yellow A,
safflower yellow B, and anhydrosafflor yellow B are the pre-
dominant yellow pigments. These pigments are glycosylated
flavonoids and thus have high water solubility in contrast to
carotenoids.
A significant variation in literature among total, free,
and esterified sterol exists. The phytosterol content in saf-
flowers is approximately 20004500 mg g 1 seed. The highest
sterol content was obtained in seeds at 24 days after anthesis
(i.e., flowering) and stabilized at 2400 mg g 1 seed at day 47
(i.e., end of maturity). Therefore, the variation in phytosterol
data may be due to seed maturity. Phytosterol contents of
15205745 mg g 1 of oil have been reported. In general, the
free (48%) sterol form accounts for the majority of the phy-
tosterol content followed by esterified (39%) form. Steryl
glycosides and acyl steryl glycosides collectively make up
about 13% of the total sterols. Regardless of the sterol form,
b-sitosterol accounts for the largest percentage (5070%) of
the phytosterols. Campesterol, D7-stigmastenol, D5-
avenasterol, and stigmasterol each make up 612% of the
total phytosterol content. Reductions in total sterols due to
maturity and refining have been documented; however, the
percentage of each phytosterol remains relatively consistent,
for example, b-sitosterol accounting for the greatest percentage.
Phenolic Components
The majority of the phenolic compounds of safflower are
located in the flower petals and meal component of the seed.
The meals contain 38005700 mg g 1 of phenolic glycosides.
Although humans find these compounds bitter, animals
appear to be less impacted by the bitterness, and thus, saf-
flower is often used as a protein source in animal rations.
Total phenolic contents of water extracts were 40 and
126.0 mg gallic acid equivalence (GAE) per gram flowers and
seed, respectively. In contrast, only 26 mg GAE per gram was
observed in the oil. Flavonoid contents of safflower water
extract obtained from the seed were 62.2  1.9 mg quercetin
equivalence per gram, while an extract from dried flowers
contained approximately 10 catechin equivalences per gram
extract.
Further characterization of the water extract of the seed
indicated that the main phenolic compound was
( )-epigallocatechin (109.6 mg g 1). Other phenolics includ-
ing 4-hydroxy-benzhydrazide derivative (18.2 mg g 1), 2-
amino-3,4-dimethylbenzoic acid (16.8 mg g 1), and gallocate-
chin (17.0 mg g 1) account for the majority of the remaining
phenolic compounds. The phenolic acids trans-ferulic,
 THE OILSEEDS | Overview of the Oilseed Safflower (Carthamus tinctorius L.)
chlorogenic, p-coumaric, and syringic acids are present at 3.0,
2.4, 0.5, and 0.2 mg g 1 extract, respectively. Additional flavo-
noids included naringin, rutin hydrate, quercetin dehydrate,
luteolin, and kaempferol at concentrations of 6.0, 3.7, 2.2, 1.6,
and 0.8 mg g 1, respectively. Chlorogenic acid, quercetin-3-
galactoside, and gallic acid are present at concentrations of
3742, 1516, and 1012.1 mg g 1 dried flower petals, respec-
tively. Other phenolic compounds were present at concentra-
tions less than 9 mg g 1. Although differences in concentrations
of phenolics were observed, the general trend in individual
phenolics remained consistent regardless of growing location
and cultivar.
Uses
Although considered a modern use, safflower as a vegetable oil
dates to around 260 BC. The use of safflower as a vegetable oil
is limited due to the availability of inexpensive commodity
oils. However, high-oleic acid safflower has promise as frying
oil due to the low level of polyunsaturated fatty acids. Many
oxidative stability index evaluations indicate that high-oleic
acid safflower oil has comparable oxidative stability to that of
high-oleic acid sunflower oil. The oxidative stability of high-
oleic acid safflower was further enhanced in a safflower line
that contained high g-tocopherol content. The non-
saponifiable fraction of safflower was found to inhibit frying
oil polymerization.
In countries such as Korea, safflower oil is used as condi-
ment oil and is therefore roasted prior to oil extraction. Mild
roasting of the safflower seed does not affect fatty acid compo-
sition, but increases in tocopherols and oxidative stability have
been documented by several researchers. However, excessive
heat treatments such as boiling and microwaving decrease
tocopherol concentration and oxidative stability. In addition
to tocopherol and Maillard browning compounds, formed
during roasting, phytosterols and phenolic compounds are
thought to stabilize the safflower oil.
Extracts of safflower have been shown to exhibit antioxi-
dant activity and thus could potentially serve as a source of
natural antioxidants. Water extracts of safflower seeds showed
dose-dependent activity in several radical scavenging assays.
The ORAC value of 63 mM Trolox equivalence per gram of
seed extract was comparable to that of tomatoes. Radical scav-
enging properties were also found for extracts of the flower
petals. Roasting increased slightly the Trolox equivalent antiox-
idant capacity (TEAC), while microwaving of the seed signifi-
cantly increased TEAC compared to the control or raw seed. The
list of studies supporting the antioxidant activity of safflower
continues to grow. The diverse phenolic composition, yellow
pigments, and tocopherol are just a few examples of compounds
that likely contribute to the antioxidant activity.
In addition to their antioxidant activity, the pigments of the
flowers have been used extensively as a food, cosmetic, and
textile dye. Hydroxyl safflower yellow A, safflower yellow B,
and anhydrosafflor yellow B are the yellow pigments of
safflower, while carthamin is the red color. Unlike the yellow
pigments, carthamin has limited water solubility and is there-
fore used in lipid-based products such as chocolate. The yellow
pigments have been used to color cheese and sausage, juice,
257
and candy. The stability of the pigments can be problematic
under the presence of light. Carthamin in particular has poor
stability when added to aqueous-based food systems. The sta-
bility improves in a 75% sucrosecholine solution compared
to water. A 90% glucosecholine chloride solution had the
greatest stabilizing effect on carthamin. Although degradation
occurred, carthamin had greater stability in a xylitolcholine
solution than in water under thermal degradation conditions.
In contrast to carthamin, the yellow pigments tend to be more
stable but tend to degrade slightly under acid conditions. Solu-
tion containing lactic acid and ethanol facilitated the rapid
degradation of carthamin and to a lesser extent the yellow
pigments. Multiple studies have demonstrated the stability
issues of safflower pigment. The importance of the food matrix
in stabilizing/destabilizing the colorants must be considered if
safflower pigments are to be used in foods.
Traditional Chinese medicine has used safflower as a cure
for many ailments. However, until relatively recently, evidence
supporting the use of safflower as a treatment for many
ailments has been lacking. A summary of the pharmacological
and medicinal benefits can be found in the literature.
However, a few examples include the antiatherogenic effect
of N-(p-coumaroyl)serotonin and antiadipogenic effects of
N-feruloylserotonin of safflower seed extract. The serotonin
derivatives are believed to inhibit the activation of the vascular
smooth muscle cells, thereby preventing atherosclerosis.
Alcohol extracts of the flowers had matrix metalloproteases-2
(MMP-2) inhibitory activity. MMP-2 are important for many
biological functions but if not properly controlled can contrib-
ute to the development of diseases such as cancer and athero-
sclerosis. Therefore, the MMP-2 inhibitory activity of hydroxyl
safflower yellow A and safflower yellow B could be important
treatments for disease affected by MMP-2.
The use as a clothing dye and a coloring for food has been
documented. Safflower as a potential source of health-
promoting oil (i.e., high g-linolenic acid) and as a biodiesel
feed stock has attracted attention from commercial entities.
Safflower is also a common bird food. Recently, pharmacolog-
ical and medicinal benefits of safflower have attracted the
attention of health researchers. Therefore, the author invites
readers to review these source materials.
Future Prospects
A significant body of knowledge exists regarding the produc-
tion and composition of safflower. However, additional pro-
duction research related to stressed environments such as high
salinity and low moisture conditions are needed, specifically
improving yield under stress conditions. Furthermore, the use
of molecular techniques to alter or enhance composition is
needed. The genetic manipulation of safflower to produce
high g-linolenic acid safflower oil represents an opportunity
to produce safflower oil with eicosapentaenoic acid and doc-
osahexaenoic acid, that is, long chain omega-3 fatty acids are
important for health and are normally found in fish and algae.
Safflower is unique among many oilseeds in that temperature
during growing and seed filling does not significantly affect
unsaturated fatty acid content. Therefore, greater production
258 THE OILSEEDS | Overview of the Oilseed Safflower (Carthamus tinctorius L.)
area could be achieved in contrast to oilseeds that have
temperature-dependent regulation of unsaturated fatty acids.
Expanding the understanding of the functional characteris-
tics of safflower seed and flower is needed. Several studies have
documented the color stability of flower extracts in various
model systems. However, additional investigation is needed in
food-based systems. The bitterness associated the meal, while
not a deterrent in animal feed, is not well suited for human
consumption. Identifying ways to reduce or eliminate the bit-
terness at a commercial level is needed. Alternatively, methods
to extract and purify the bitter compounds could be justified if
sufficient bioactives could be isolated. Some of the bitter com-
pounds have biological activity in model systems. Traditional
Chinese medicine has used safflower for treatment of a number
of aliments; however, the active components are not fully char-
acterized. Although a number of studies have been completed,
there is still a need to assess the bioactivity of safflower.
Exercises for Revision
What parts of the safflower can be used for food, feed,
biofuel, coloring, and bioactive compounds?
Why is safflower grown worldwide?
What factors influence the growth of safflower?
Why is the taproot of safflower important?
How does growing environment affect safflower seed
composition?
Why is safflower unique among oilseeds in terms of fatty
acid composition?
What bioactive compounds are present in safflower and
what part of the plant can these compounds be found?
What type of safflower oil can be used as frying oil? What
makes this oil well suited for frying applications?
Why is safflower a potential source of omega-3 lipid?
Why is the yellow pigment not commonly used in food
systems?
Exercises for Readers to Explore the Topic Further
How does safflower differ from other oilseeds in terms of
production and oil content and fatty acid composition?
Cross-reference to oilseed articles.
Identify several countries that produce safflower and com-
pare production practices and oilseed qualities among the
produced seeds? Do any countries use wild safflower?
Why is defatted safflower meal used only for livestock feed
and not human food? How does this compare to other
oilseed meals? Cross-reference to oilseed articles.
See also: Bioactives and Toxins: Bioactives: Antioxidants; Fats:
Healthy Fats and Oils; Lipid Chemistry; Food Grains and Well-
being: Functional Foods: Overview; Genetics of Grains:
Development of Genetically Modified Grains; The Oilseeds: Canola:
Overview; Linseed: Overview; Oilseeds: Overview; Soybean: Overview;
Sunflower: Overview; Cottonseed: Overview.
Further Reading
Asgarpanah J and Kazemivash N (2013) Phytochemistry, pharmacology and medicinal
properties of Carthamus tinctorius L. Chinese Journal of Integrative Medicine 19(2):
153159.
Esteban A, Sicardo M, Mancha M, and Martnez-Rivas J (2004) Growth temperature
control of the linoleic acid content in safflower (Carthamus tinctorius) seed oil.
Journal of Agricultural and Food Chemistry 52: 332336.
Hamrouni-Sellami I, Salah H, Kchouk M, and Marzouk B (2007) Variations in
phytosterol composition during the ripening of Tunisian safflower Carthamus
tinctorius L. seeds. Pakistan Journal of Biological Sciences 10(21):
38293834.
Kisha T and Johnson R (2012) Safflower. In: Gupta SK (ed.) Technological Innovations
in Major World Oil Crops. Breeding, Vol. 1, pp. 147164. Berlin: Springer.
Kizil S, Cakmak O, Kirici S, and Inan M (2008) A comprehensive study on safflower
(Carthamus tinctorius L.) in semi-arid conditions. Biotechnology and
Biotechnological Equipment 22(4): 947953.
Li B, Zong X, Zhang H, et al. (2014) Screening and structural characterization of mmp-2
inhibitor from Carthamus tinctorius L. using ultrafiltration liquid
chromatographymass spectrometry. Journal of Liquid Chromatography and
Related Technologies 37: 23272336.
Luterottia S, Franko M, Sikovec M, and Bicanic D (2002) Ultrasensitive assays of trans-
and cis-b-carotenes in vegetable oils by high-performance liquid
chromatographythermal lens detection. Analytica Chimica Acta 460: 193200.
Mariod A, Ahmed S, Abdelwahab S, et al. (2012) Effects of roasting and boiling on the
chemical composition, amino acids and oil stability of safflower seeds. International
Journal of Food Science and Technology 47: 17371743.
Mayerhofer M, Mayerhofer R, Topinka D, Christianson J, and Good A (2011)
Introgression potential between safflower (Carthamus tinctorius) and wild relatives
of genus Carthamus. BMC Plant Biology 11: 47. http://dx.doi.org/10.1186/1471-
2229-11-47.
Nykiforuk C, Shewmaker C, Harry I, et al. (2012) High level accumulation of gamma
linolenic acid (C18:3D6.9,12 cis) in transgenic safflower (Carthamus tinctorius)
seeds. Transgenic Research 21: 367381.
Ortega-Garca J, Gamez-Meza N, Noriega-Rodriguez J, et al. (2006) Refining of high
oleic safflower oil: Effect on the sterols and tocopherols content. European Food
Research and Technology 223: 775779.
Singh V and Nimbkar N (2006) Safflower (Carthamus tinctorius L.). In: Singh RJ (ed.)
Genetic Resources, Chromosome Engineering, and Crop Improvement, Vol. 4.
Oilseed Crops, pp. 167194. Boca Raton, FL: CRC Press, Taylor & Francis Group.
Smith J (1996) Safflower. Champaign, IL: AOCS Press.
Vilatersana R, Garnatje T, Susanna A, and Garcia-Jacas N (2005) Taxonomic problems
in Carthamus (Asteraceae): RAPD markers and sectional classification. Botanical
Journal of the Linnean Society 147: 375383.
Weiss EA (2000) Safflower. In: Weiss EA (ed.) Oilseed Crops, pp. 93129. Oxford:
Blackwell Science.
 Linseed: Overview
S Cloutier, Agriculture and Agri-Food Canada, Ottawa, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Linseed is an ancient crop that has been useful and versatile
through the ages.
Linseed is a minor oilseed crop with a unique fatty acid
profile.
Linseed oil is rich in alpha-linolenic acid (ALA) that gives
the oil unique drying properties.
Many health benefits have been attributed to the consump-
tion of linseed.
Low yield relative to other major oilseeds limits the expan-
sion of the crop.
Learning Objectives
In the following article, the reader is expected to understand
the uniqueness of linseed compared to other oilseed crops.
An overview of the agronomic practices and production
limitations is presented to illustrate the challenges associ-
ated with the development of improved cultivars and of
cultivating this crop. Emphases have been placed on not
only the composition of linseed as it relates to its multiple
end uses but also the multiple health benefits associated
with the consumption of linseed.
Introduction
The species epithet of the Latin name of flax, Linum usitatissi-
mum L., means most useful, an appropriate name because this
crop has been highly versatile and esteemed through the ages.
Flax has two morphotypes: fiber and oilseed. Tall and
branched only in the top quarter of the plant, fiber flax is
grown for its stem fibers that are extracted and processed into
linen, nonwoven mats, paper, and various other products.
Oilseed flax plants, also called linseed or flaxseed, are shorter
and more branched and have larger seeds than the fiber type.
Flax is one of the worlds oldest cultivated crops, praised by
ancient civilizations for its fiber and seed uses. Today, varieties
are grown for either stem fiber or seed, although value can be
extracted from the seeds of fiber flax and, conversely, the stem
fiber of linseed. Over the last 50 years, the worlds production
of linseed fluctuated between 2 and 4 million tonnes (Mt), but
in the last decade, it has stabilized at a little more than 2 Mt
with the largest producers being Canada, the Russian
Federation, and China. In 2012, world oil production from
the top 12 major oilseed crops amounted to 157 Mt, of which
linseed oil production at 544 t accounted for a mere 0.35%. As
such, linseed is a minor oilseed crop worldwide, but the
unique characteristics of its oil combined with its high content
of seed dietary fibers and lignans respond to specific markets
not addressed by the major oilseeds.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00031-0
The term linseed often refers to the seed destined for oil
extraction with industrial end uses while the term flaxseed is
more commonly associated with nonprocessed seed used
directly in human food or animal feed. In reality, the two
terms represent flaxseed with the same oilseed quality and
they can be used interchangeably. Currently cultivated linseed
varieties contain 4050% oil, of which 5557% is ALA, an
essential fatty acid. LinolaTM is a trademark name of the
solin (generic term) type of flax that contains < 5% linolenic
acid, resulting in oil similar in composition to sunflower. In
Canada, LinolaTM or solin varieties were required to have a
yellow seed coat to make them visually distinguishable from
brown seeded linseed varieties, although there are also yellow
linseed varieties. High linolenic acid content varieties charac-
terized by more than 60% ALA are generically referred to as
high-lin such as the European variety Omega or under the
registered name Nulin.
End Uses
An overview of the end use versatility of fiber flax and linseed is
illustrated (Figure 1). For the purpose of this article, the focus
will be strictly on the seeds. Linseed is used as human food in
baked goods and other products, either whole or ground.
Whole seeds can also be fed to poultry because they can
grind the seeds with their gizzard. The processing of linseed
to extract oil produces a by-product called meal. Rich in pro-
tein and energy, the meal is a high-value feed for livestock. The
extracted oil can be utilized as human food, mostly as a sup-
plement, but its main application is as industrial oil in, for
example, stains, paints and linoleum flooring.
Composition
Expressed on a dry basis, typical linseed contains 4143% oil,

28% dietary fiber, 2123% protein, 6% carbohydrate, and
4% ash but these can vary substantially among genotypes,
particularly oil content. In genebanks, oil contents ranging
from 31.4% to 45.7% were reported while levels up to 52.3%
were recorded in field plot trials. Most modern linseed cultivars
field-grown in temperate climates contain 4549% oil.
The fatty acid composition of linseed oil is unique among
plants because it is particularly high in ALA, the parent fatty
acid of the omega-3 family. The major oilseeds of the world
contain much less ALA than linseed (Figure 2). Rapeseed
(canola) and soybean oils have
10% and 8% ALA, respec-
tively, while sunflower, cottonseed, corn, and peanut oils only
have traces (<1%). Conventional linseed varieties typically
contain 57% ALA, 16% linoleic acid (LA), 18% oleic acid, 4%
stearic acid, and 5% palmitic acid. With 73% polyunsaturated
fatty acid content, linseed oil has unique drying properties.
259
260 THE OILSEEDS | Linseed: Overview
Extraction
Oil Meal Processing
Animal
Food Industrial
feed
Linoleum Linen, paper,
coatings, paints, nonwoven mats,
stains, insulation, fire
lubricants, etc. logs, etc.
Figure 1 The versatility of flax (Linum usitatissimum L.). Stem fibers
extracted from the fiber morphotype can be processed into high-quality
linen. Lower-quality fibers or other straw components such as the
shives can be used to produce a wide variety of products. Whole seeds
from the oilseed morphotype (linseed or flaxseed) can be used in the
food industry or as animal feed. Linseed oil extraction generates a
by-product called meal, which is an excellent animal high-protein feed
source. The highly unsaturated oil has some food usage in salad dressing
or as a functional food, but its tendency to quickly become rancid
makes special handling a requirement. The uniqueness of the fatty acid
composition of linseed oil gives it distinctive drying properties
desirable for many industrial applications.
LinolaTM or solin varieties, developed through mutation breed-
ing and selected for their inactive fatty acid desaturases 3A
and 3B, have < 5% ALA, often only 23%, but contain
6575% LA. The fatty acid composition of LinolaTM or solin
varieties is similar to conventional sunflower (Figure 2). These
varieties were developed to meet the market need of the mar-
garine industry. Unfortunately for LinolaTM, the margarine
sector developed different formulations using mostly non-
hydrogenated soybean and canola oils and the demand for
LinolaTM oil was not sustained. Today, these types of linseed
varieties are no longer commercially grown. Linseed varieties
with very high ALA content (high-lin) ranging from 60% to
75% have been developed and marketed. Similar to chia seeds
(5269%) in terms of ALA content, both conventional linseed
and high-lin varieties constitute a richer source of ALA because
their oil content of 4050% exceeds that of chia (Salvia hispa-
nica L.) seeds, which ranges from 26% to 38%. High-lin vari-
eties were developed for human food markets including the
nutraceutical and functional food sectors, but they can also be
used as a high ALA source for blending of industrial linseed oil
to ensure the desired oil properties.
0% 20% 40% 60% 80% 100%
Linseed
Rapeseed
Soybean
Solin
Sunflower
Cottonseed
Corn
Peanut
Saturated fat Monounsaturated fat
Linoleic acid Alpha linolenic acid
Figure 2 Fatty acid composition of linseed and solin (LinolaTM) oil
compared to six of the major oilseed crops of the world. With
57%
alpha-linolenic acid (ALA) content, linseed oil is unique because none of
the other major oilseed crops exceed 10% ALA. Solin (LinolaTM) are
varieties selected through mutation breeding that have <5% ALA
content, and as such, their overall fatty acid composition is similar to
sunflower oil. Data obtained from Souci SW, Fachmann W, and Kraut H
(2008) Food Composition and Nutrition Tables, 7th edn. Boca Raton, FL:
CRC Press, Taylor & Francis Group.
Linseed contains 2123% protein on a dry-weight basis but
the meal obtained after oil extraction has an enriched protein
fraction of 3133%. The amino acid composition is considered
to be generally good because it contains all of the essential
amino acids.
On a dry-weight basis, linseed has
28% dietary fiber with
a soluble to insoluble fiber ratio ranging from 20:80 to 40:60.
These dietary fibers (not to be confused with the stem fibers)
are for the most part present in the seed coat. In a small subset
of varieties, mucilage (main soluble fiber of linseed) content
was shown to vary from 3% to 8% of the dry seed weight
indicating that the variation for this trait would possibly be
more substantial in a diverse panel of accessions. Mucilage
dissolved in water is viscous and this viscosity can be a quan-
titative measurement of its quality. In a study of Canadian
varieties, the mean mucilage indicator values ranged from 90
to 246 cSt ml g1. Hence, two varieties with the same mucilage
content may have very different functional properties.
Health Benefits
Both ALA and LA are essential fatty acids because humans do
not have the ability to synthesize them from their precursors.
We must incorporate them into our diet because they play
important physiological roles. In the human body, they can
be further elongated and desaturated to produce docosahexae-
noic acid (DHA), eicosapentaenoic acid (EPA), arachidonic
acid (AA), and other very long polyunsaturated fatty acids
with proven functionalities. ALA, DHA, and EPA are omega-3
fatty acids whose health benefits include roles in reduction of
blood pressure, blood triglycerides, inflammation, and the

incidence of cardiovascular diseases. Recently, Health Canada
has granted a health claim regarding the effectiveness of
ground flaxseed on lowering cholesterol allowing health
claims such as the following to be made on flaxseed labels:
16 g or 2 tablespoons of ground flaxseed supplies 40% of the
daily amount shown to help lower cholesterol. On the other
hand, LA can be converted into omega-6 AA, which has been
shown to be detrimental by promoting inflammatory reac-
tions. The enzymes delta-6, delta-5, and delta-4 desaturases
are shared between the omega-3 and omega-6 pathways. For-
tunately, the high omega-3 to omega-6 ratio of linseed favors
the omega-3 pathway over the omega-6 pathway. The Western
diet is generally considered to have an omega-6 to omega-3
ratio that is too high (20:1) due to the types of fats and oils
consumed (e.g., soybean; Figure 2) and hence favoring the
omega-6 pathway. The replacement of some of these fats and
oils with whole and ground linseed or linseed oil would rep-
resent a healthy alternative to meet the recommended 2:1 to
6:1 ratio.
Linseed contains both soluble and insoluble dietary fibers.
The predominant fraction of the soluble fiber is the mucilage,
which has been associated with lowering of blood cholesterol
and glucose. Lignans are phytoestrogens that are also part of
the dietary fiber component. Linseed is one of the best
sources of these diphenolic compounds. Secoisolariciresinol
diglucoside, the major linseed lignan, has been associated
with reduced risks of hormone-sensitive cancers such as pros-
tate, colon, and breast cancers. Lignans also have antioxidant
properties leading to their positive role in reducing the activity
of free radicals.
Antinutritional Factors
There are two main well-documented antinutritional compo-
nents in linseed: cyanogenic glucosides and linatine. The two
main cyanogenic glucosides contained in mature seeds are
linustatin and neolinustatin. Following ingestion, hydrolysis
can lead to the release of hydrogen cyanide that can impede
the essential process of cell respiration. Fortunately, the
amount of cyanogenic glucosides in linseed is low, and
because the acute lethal dose of hydrogen cyanide in human
is 0.53.5 mg Kg1 of body weight, an average adult would
have to consume
1000 g of linseed in a day to receive a fatal
dose. The recommended daily intake of linseed of 3040
grams contains well below the toxic dosage. Moreover, the
acidic environment of the human stomach is not conducive
to cyanogenic glucoside hydrolysis. As a consequence, these
toxic degradation products are more of a concern in
ruminants.
Linatine is an antagonist of vitamin B6. Symptoms associ-
ated with B6 deficiencies were observed in young poultry;
hence, their ration should be limited to <10% linseed meal.
As mentioned earlier, the mucilage comprises soluble dietary
fibers well known for their positive role in laxation. In animal
feed, the outcome of rapid passage in the intestinal tract is poor
nutrient absorption and its consequent performance reduc-
tion, particularly in poultry, a consequence that also limits
the linseed meal proportion of the ration.
THE OILSEEDS | Linseed: Overview 261
Animal Feed and Omega-3-Enriched Products
Linseed meal, ground or whole flaxseed, can be used as feed for
poultry, swine, and cattle because it is a good source of protein,
oil, and energy although whole flaxseed can only be used with
poultry. There are several reports on the health benefits asso-
ciated with feeding linseed-derived products to livestock. How-
ever, as outlined in the earlier-mentioned section, livestock
producers must follow recommended guidelines to avoid the
side effects associated with deficiencies, antinutritional effects,
or ability to process the ration.
A main driver for using linseed as feed is for the production
of omega-3-enriched eggs and meat. The feeding of 10% lin-
seed ration to laying hens increased the omega-3 content in
eggs by a factor of 4. In Canada, a minimum of 300 mg of
omega-3 fatty acid per serving is required to allow the labeling
of a product as a source of omega-3. Such labeling commands a
premium that health-conscious consumers are willing to pay.
Omega-3-enriched chicken and pork products are also avail-
able to consumers.
Agronomy
In temperate climates, linseed is grown as a summer annual
crop that matures in 90150 days including a vegetative, a
flowering, and a seed maturation period. In some areas of the
world, winter-type linseed is grown. These varieties do not
require vernalization per se to flower but they have the ability
to endure some cold temperatures in the early developmental
stages. While each boll may contain a maximum of ten seeds,
linseed and fiber flax accessions in western Canadian field
trials produced only 3.58.1 seeds per boll. The flat, oval,
and pointed seeds are small with thousand-seed weight rang-
ing from 2.7 to 8.4 g, but the majority of Canadian and US
linseed varieties have thousand-seed weight of 55.5 g.
Ideally, linseed is sown in firm, moist soil 1.53.0 cm deep
in rows 1520 cm apart at a seeding rate of 3045 kg ha1.
Early seeding is recommended to avoid some diseases (see
succeeding text) and drought and heat during flowering and
seed filling, which translates into higher yield, oil content, and
quality. Heat and drought stresses at flowering time and during
seed filling result in significant yield losses and lower oil con-
tent. Seeding practices for linseed differ greatly from those of
fiber flax where a high plant density is critical to fiber produc-
tion and quality. In general, the utilization of agronomic best
management practices such as early seeding, crop rotation, and
weed and disease control has a major effect on linseed yield.
Genetics
Linum usitatissimum L. is a diploid species with 15 pairs of
chromosomes (2n 30). Its genome is composed of an esti-
mated 373 Mbp of nucleotides for which an assembled
sequence is available (www.phytozome.net). There are 48 000
accessions of cultivated flax preserved in genebanks around the
world but only 10 000 are estimated to be unique because of
the redundancy between collections. Nevertheless, these
262 THE OILSEEDS | Linseed: Overview

collections represent a tremendous amount of diversity for
genetic enhancement (Figure 3). Other important genetic
resources include a large collection of expressed sequences
(genes), a consensus genetic map, and a physical map that
were used to order and annotate the genome sequence.
A core collection of flax including more than 300 linseed
types and 92 fiber types was extensively characterized in the
field at two locations during four consecutive years. Agronomic
traits such as yield and its components (thousand-seed weight,
seeds per boll, and bolls per area), lodging, maturity, flowering

Figure 3 Genetic diversity in cultivated flax, Linum usitatissimum. (a) Flower color varies from white to shades of pink or blue and petals overlap,
margin, and folding are also indicative of the breadth of the genetic diversity. (b) At maturity, some accessions have completely closed bolls (or
capsules) while others are slightly or completely dehiscent. The latter is a useful trait to disseminate the seeds in the wild but is undesirable and selected
against by breeders for usage as a crop. (c) The seed coat color of linseed can vary from bright yellow to olive to dark brown and even include variegated.
Photos, reproduced with permission, are courtesy of Dr. Axel Diederichsen, Plant Gene Resources of Canada, Saskatoon, SK, Canada.

time, height, and plant branching were measured. Seed quality
traits such as fatty acid composition, iodine value, oil content,
and protein content were also evaluated. Finally, disease rat-
ings for rust, pasmo, and powdery mildew were recorded. The
collection was genotyped with molecular markers and a
genome-wide association study revealed several marker-trait
associations. These important studies provided background
information on the flax genome and the genetics of important
traits that, combined together, will assist and accelerate
improvement of linseed by plant breeders.
Yield
In western Canadian field plots, yield of linseed accessions as
high as 2.5 tonnes per hectare (t ha1) was reported, but higher
yields have been measured (Paul Dribnenki, personal
communication, January 2014). However, this yield potential
is not achieved for the crop at large and the yields of linseed
worldwide remain at < 1 t ha1. Even in the main producing
countries, it is still only 1.4 t ha1 and the rate of yield increase
remains below 1.5% per year, far less than its oilseed compet-
itors canola (1.62 t ha1) and soybean (1.62.6 t ha1). Yield
improvement is therefore the main objective of linseed breed-
ing programs because it is the single most important trait to
ensure competitiveness. In recent years, plant breeders have
realized that the narrow genetic base of the germplasm has
limited yield improvement. To overcome this limitation, lin-
seed breeders are now incorporating exotic material, using
modified breeding schemes, and applying molecular markers
into breeding programs. For example, backcrosses of molecular
markers associated with thousand-seed weight to introgress the
trait into adapted elite germplasm are being implemented.
Genomic selection could also prove useful toward yield
improvement, although there are no reports yet on its usage
in linseed improvement.
Many oilseed crops such as corn, canola, and soybean have
well-established hybrid systems that can take advantage of
heterosis, also known as hybrid vigor. As mentioned earlier,
linseed is highly self-pollinated, and to date, a single hybrid
has been released in China. Although hybrid production of
linseed bears promise and is theoretically possible, the devel-
opment of an economically viable and stable system poses
major challenges.
Diseases
Several diseases can affect the crop during its growing season.
The most important are fungal diseases that are prevalent in
most growing areas of the world: fusarium wilt caused by
Fusarium oxysporum, rust caused by Melampsora lini, pasmo
caused by Septoria linicola, and powdery mildew caused by
Oidium lini. The crop can also be affected by other fungal,
phytoplasma, and viral diseases but these tend to either be
localized or have a negligible impact on yield or quality.
Fusarium wilt, a soil-borne pathogen, can wreak havoc on a
crop because heavy infection of susceptible varieties results in
complete wilting of the plants and major yield reduction. The
combination of moderate resistant varieties and crop rotation
THE OILSEEDS | Linseed: Overview 263
to keep the soil inoculum low are effective ways of controlling
the disease. The host disease resistance mechanism seems to be
controlled by a few major genes meaning that the development
of resistant varieties should be highly feasible in a reasonable
time frame. However, the large environmental component
involved in the expression of wilt tolerance challenges plant
breeders and continues to impede breeding progress.
Rust spores are airborne and can infect flax plants when
temperature and moisture conditions are conducive. Visual
symptoms are the presence of pustules with or without chlo-
rosis or necrosis on the aerial parts of the plants. Interestingly,
it was while studying flax rust in the 1950s that Harold Henry
Flor came up with the gene-for-gene concept, stating that for
each gene conditioning rust reaction in the host, there is a gene
counterpart in the fungus conditioning pathogenicity. This
concept has since been found to be applicable to many plant
diseases. In flax, several rust resistance genes (R genes) have
been cloned in the last two decades assisting in their introgres-
sion in breeding lines, an important asset to breeders because
rust-resistant varieties are the best way to counter this disease.
Powdery mildew infection appears as a white mass of myce-
lium and spores on the leaves. Heavy infection can occur under
high humidity conditions. Similarly to fusarium wilt, resis-
tance seems to be controlled by a few major genes. As with
the other two previous fungal diseases, genetic resistance exists
and planting resistant cultivars is the best practice to control
the disease in areas where it is prevalent.
Pasmo is the most prevalent disease on the Canadian Prai-
ries. Early infection of pasmo has a major detrimental impact
on yield and quality. The early brown spots can evolve into
complete leaf desiccation. The disease girdles the stems and
increases the prevalence of lodging, which has also been shown
to cause significant yield reduction in flax and negatively affect
seed quality. Early seeding of clean seeds, crop rotation, and
fungicide application are the best control methods because
commercial cultivars released to date seem to lack resistance
to this disease.
Transgenic
In 1996, CDC Triffid was the first and only genetically modi-
fied (GM) linseed variety to be registered in Canada. CDC
Triffid was the result of the transformation of NorLin with a
modified Arabidopsis acetolactate synthase gene that conferred
resistance to residual soil activity of sulfonylurea herbicides.
Breeder seed of the variety was distributed to seed growers for
pedigreed seed production. In 199798, in light of the Euro-
pean Union (EU) moratorium on GM crops and foods, the
seed company owning the right to CDC Triffid voluntarily
halted its seed multiplication and coordinated its retrieval
and destruction with the Flax Council of Canada to prevent
potential harm to the industry. CDC Triffid pedigreed seed
production was recalled and destroyed. In 2009, a shipment
of linseed from Canada to the EU tested positive for a well-
known GM marker. Further testing confirmed the low-level
presence of CDC Triffid. While this variety had received
approval for feed and industrial uses in Canada and the United
States, it constituted a nonapproved event in the EU and there
is zero tolerance for such. Canada and the EU entered into
264 THE OILSEEDS | Linseed: Overview
discussions to mitigate the crisis that led to the establishment
of a stewardship program and testing protocols. In 2013, one
can conclude that the monitoring has been a success because
the low-level presence of the CDC Triffid gene construct in
Canadian linseed has been substantially reduced. Unfortu-
nately, exports to the EU are only half what they were prior
to 2009 indicating that market confidence remains to be fully
regained.
Conclusion
Linseed is a unique oilseed crop that combines high oil content
with high content of ALA giving the oil distinctive drying
properties sought in the manufacturing of biodegradable
industrial products. ALA, lignans, and high fiber content of
linseed are also contributing to making linseed highly suitable
for the food and feed markets because these seed components
have been linked to several health attributes in human and
animals. While linseed remains a minor oilseed crop world-
wide, its versatility and uniqueness continue to be unmatched
by the main oilseed crops.
Exercises for Revision
What components of linseed are associated with health
attributes?
What is the main fatty acid of linseed and how does its
composition compare to other major oilseed crops?
What makes ALA an essential fatty acid?
Why is Linum usitatissimum L. considered a dual purpose or
total utilization crop?
What morphological features distinguish fiber flax from
oilseed flax (linseed)?
What is the main characteristic distinguishing the solin-type
(LinolaTM) from the conventional linseed?
How is linseed used to create omega-3-enriched chicken
and pork meats?
What are some of the main production challenges currently
limiting the expansion of linseed as a crop?
Exercises for Readers to Explore the Topic Further
Molecular genetics is making great strides in gene discovery
for discrete and quantitative traits but the translation of this
research into breeding is generally slower in minor crops
such as linseed because of limited investments. Phenotyp-
ing is expensive but necessary to identify and validate func-
tional genetic variants. To survive and thrive as a crop,
linseed yield must be improved and further research toward
that goal will certainly encompass the genomics
translationphenotyping continuum including also agron-
omy and disease research activities. Genomic selection, a
method of predicting breeding values using genomic
sequencing, is proving viable in many major crops such as
wheat, corn, barley, and rice and bears promise for other
crops such as linseed that has the advantage of a small
genome. Genetic transformation has the potential to bring
about novel products because linseed is readily amenable
to this type of technologies. As a health food, the anti-
nutritional compounds of linseed remain and will continue
to be an important research target and the topic of various
research activities.
Acknowledgments
The author sincerely thanks Drs. Helen Booker, Axel
Diederichsen, Paul Dribnenki, and Gordon Rowland and
Mr. Andrzej Walichnowski for their comments and edits
made to this article. The author also thanks Ms. Michelle Beaith
and Drs. Allan Green and Paul Dribnenki for the information
on the status of LinolaTM around the world and Ms. Kelley
Fitzpatrick for the information on the regulatory affairs of
flax health claims.
Further Reading
Anonymous (2008) Flax feed industry guide. Winnipeg: Flax Canada 2015.
Diederichsen A (2007) Ex situ collections of cultivated flax (Linum usitatissimum L.)
and other species of the genus Linum L. Genetic Resources and Crop Evolution
54: 661678.
Diederichsen A, Kusters PM, Kessler D, Bainas Z, and Gugel RK (2013) Assembling
a core collection from the flax world collection maintained by Plant Gene Resources
of Canada. Genetic Resources and Crops Evolution 60: 14791485.
Dribnenki JCP and Green AG (1995) Linola 947 low linolenic acid flax. Canadian
Journal of Plant Science 75: 201202.
FAOSTAT (2013) Production: Crops. Available at http://faostat3.fao.org/home/index.
html (accessed December 2013).
Fofana B, Ragupathy R, and Cloutier S (2010) Flax lipids: Classes, biosynthesis,
genetics and the promise of applied genomics for understanding and altering of fatty
acids. In: Gilmore PL (ed.) Lipids: Categories, Biological Functions and
Metabolism, Nutrition and Health, pp. 7198. New York: Nova Science
Publishers Inc.
McHughen A, Rowland GG, Holm FA, Bhatty RS, and Kenaschuk EO (1997) CDC Triffid
transgenic flax. Canadian Journal of Plant Science 77: 641643.
Muir AD and Westcott ND (2003) Flax: The genus linum. Boca Raton, FL: CRC Press.
Rashid KY (2003) Diseases of flax. In: Bailey KL, Gossen BD, Gugel RK, and
Morrall RAA (eds.) Diseases of Field Crops in Canada, 3rd edn., pp. 147154.
Saskatoon: University Extension Press.
Ryan CD and Smyth SJ (2012) Economic implications of low-level presence in a
zero-tolerance European import market: The case of Canadian Triffid flax.
AgBioforum 15: 2130.
Soto-Cerda BJ, Duguid S, Booker H, Rowland G, Diederichsen A, and Cloutier S
(2014a) Association mapping of seed quality traits using the Canadian flax (Linum
usitatissimum L.) core collection. Theoretical and Applied Genetics 127: 881896.
Soto-Cerda BJ, Duguid S, Booker H, Rowland G, Diederichsen A, and Cloutier S
(2014b) Genomic regions underlying agronomic traits in linseed (Linum
usitatissimum L.) as revealed by association mapping. Journal of Integrative Plant
Biology 56: 7587.
Souci SW, Fachmann W, and Kraut H (2008) Food composition and nutrition tables,
7th edn. Boca Raton, FL: CRC Press, Taylor & Francis Group.
Wang Z, Hobson N, Galindo L, et al. (2012) The genome of flax (Linum usitatissimum)
assembled de novo from short shotgun sequence reads. Plant Journal 72: 461473.
 The Legumes and Pseudocereals

Contents
Grain Legumes and Their Dietary Impact: Overview
Pseudocereals: Overview
Lupin: Overview
Amaranth: Overview
Beans: Overview
Buckwheat: Overview
Chickpea: Overview
Pea: Overview
Peanuts: Overview
Quinoa: Overview

Grain Legumes and Their Dietary Impact: Overview

TE Michaels, University of Minnesota, St. Paul, MN, USA
2016 Elsevier Ltd. All rights reserved.

Topic Highlights Introduction

Grain legumes have seeds that are prepared for use as

human food.
Pulse seeds are a rich source of protein in diets around the
world.
Legumes support Rhizobium bacteria that fix atmospheric
nitrogen for use by the plant and so are indispensible in
sustainable cropping systems.
Pulse consumption is constrained by low digestibility, anti-
nutritional factors, flatulence, and stigmatization as a food
for the poor.
Combining cereal and legume grains at meals provides
inexpensive plant-based diets with balanced amino acid
composition.
Seed yield can be hampered by the plants sensitivities to
environmental stress.
Grain legumes can relieve malnutrition in food-deficit
regions and protect against diet-related diseases such as
obesity, diabetes, and heart disease in food surplus regions.
Learning Objective
To obtain a comprehensive understanding of the agron-
omy, consumption, composition, antinutritional factors,
and protein complementation of grain legumes
As the field is now prepared, we shall proceed to speak of the nature
of the various kinds of grain; we must premise, however, that there
are two principal classes of grain, the cereals, comprising wheat and
barley, and the legumina, such as the bean and the chick-pea, for
instance. The difference between these two classes is too well known
to require any further description.
The Natural History. Pliny the Elder. John Bostock, M.D., F.R.S.
H.T. Riley, Esq., B.A. London. Taylor and Francis, Red Lion Court,
Fleet Street. 1855. Book 18 Chapter 9
Despite Pliny the Elders assertion that the difference
between classes of grain is too well known to warrant further
description, let us do so anyway. In this overview, we will
consider grain legumes to be plants with two major character-
istics: they belong to the legume family (Fabaceae or Legumi-
nosae) and preparations based on their dry seeds are
consumed by humans for food (although they may have
other uses as well). In contrast to cereal grains, humans partic-
ularly value legume grains as a concentrated source of protein
for our diets. Grain legumes are commonly subdivided into
partially overlapping groups according to their dominant use
and seed composition. Pulses, such as the common bean (Pha-
seolus vulgaris) and cowpea (Vigna unguiculata), are classic grain
legumes in that their dry seed composition includes high pro-
tein (>19%), high carbohydrate (>60%), and low lipid (<6%)
and where the primary culinary interest focuses on the dry seeds

Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00040-1 265

266 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview

Table 1 Names and regions of greatest genetic diversity for major grain legume species

Botanical name Common and vernacular names Center of domestication

Arachis hypogaea Peanut, groundnut Southern Bolivia, northwest Argentina

Cajanus cajan Pigeon pea, Angola pea, Congo pea, dhal, no-eye pea, red gram India
Cicer arietinum Chickpea, garbanzo bean, gram Southeast Turkey
Glycine max Soybean, soyabean, sojabean China
Lathyrus sativus Grass pea, chickling vetch Southern Europe, Southwest Asia
Lens culinaris Lentil, split pea, red dhal, masur, lenteja, lentille, manssor, burssum Eastern Mediterranean, the Fertile Crescent
(Iraq and Iran)
Lupinus albus; L. Mediterranean white lupin, narrow-leafed lupin, Australian sweet Greece, western Turkey, Eastern
angustifolius; L. mutabilis lupin, Andean pearl lupin, tarwi Mediterranean, Peru (L. mutabilis)
Macrotyloma uniflorum Horse gram Southern India
Phaseolus vulgaris Common bean, dry bean, kidney bean, frijol, caraota, poroto, Mexico and Peru
habichuela, haricot bean, snap bean
Phaseolus lunatus Lima bean Peru, Central America, the Caribbean
Pisum sativum Field pea, garden pea, guisante, muttar, pois, arveja, Alaska pea The Fertile Crescent (Iraq and Iran), Turkey,
Greece
Vicia faba Broad bean, faba bean, horse bean, Windsor bean, haba, feve Eastern Mediterranean, West Asia
Vigna radiata Green gram, golden gram, mung bean, Oregon pea, chickasano pea India, Southeast Asia
Vigna mungo Mung bean, black gram, urd, kambulu, uride India
Vigna unguiculata Cowpea, catjang, Hindu cowpea, kaffir bean, black-eyed pea, frijol, Sub-Saharan Africa
caupi

Adapted from Sinha SK (1977) Food Legumes: Distribution, Adaptability and Biology of Yield. Rome, Italy: Food and Agriculture Organization of the United Nations. Based on Sinha
(FAO) and Adams MW and Pipoly JJ III (1980) Biological structure, classification and distribution of economic legumes. In: Summerfield RJ and Bunting AH (eds.) Advances in
Legume Science, pp. 116. Kew, UK: Royal Botanic Gardens; Zohary D, Hopf M, and Weiss E (2012) Domestication of Plants in the Old World, pp. 98100. Oxford, UK: Oxford
University Press; Grabiele M, Chalup L, Robledo G, and Seijo G (2012) Genetic and geographic origin of domesticated peanut as evidenced by 5S rDNA and chloroplast; DNA
sequences. Plant Systematics and Evolution 298: 11511165.

Table 2 Protein and other nutritional constituents of grain legumes rice). Oilseed legumes, including soybean (Glycine max) and
and other reference foods (g constituent 100 g food1) peanut (groundnut, Arachis hypogaea), in contrast to pulses,
accumulate higher protein, lower carbohydrate, and higher
Food Protein Lipid Carbohydrate Water lipid. Because of the high lipid content, they are often utilized
as a source of vegetable oil for culinary and industrial uses, but
Pulses
they are also important as pulse-like ingredients in prepared
Common bean 22.2 1.1 61.5 11.5
Lima bean 21.5 0.7 63.4 10.7 foods. For example, soybean is the base of many nonfermented
Chickpea 19.3 6.0 60.7 11.5 and fermented foods such as tofu and tempeh, while peanut is
Green gram 23.9 1.1 62.6 9.0 the main ingredient in some snacks and sandwich spreads. Lupin
Cowpea 23.8 2.1 59.6 11.1 (Lupinus L.) seeds protein, lipid, and carbohydrate composition
Lupin 36.7 11.5 45.4 is intermediate to that of pulses and oilseeds, and its dominant
Oilseeds use in many areas of the world is as an animal feed. However, in
Soybean 36.5 19.9 30.2 8.5 Europe and South America, it has a long, pulse-like history of
Peanut 25.8 49.2 16.1 6.5 human consumption and now enjoys increasing popularity as a
Cereals
food crop in North America and Australia.
Wheat, hard white 11.3 1.7 75.9 9.6
Grain legumes are ancient foods that help fulfill the ageless
Brown rice, raw 7.5 2.7 76.2 12.4
Animal products need for human sustenance. The word pulse arrived in mod-
Beef, lean ground 20.0 10.0 0.0 69.5 ern English from the Greek poltos meaning porridge of meal,
Pork, ground 16.9 21.2 0.0 61.0 by way of Latin (puls and pultis), Old French (pols), and
Fish, cod 17.8 0.7 0.0 81.2 Middle English (puls).
Milk, cows 3.25% fat 3.3 3.3 4.7 88.0 Gathering plant parts for food is a reliable way (in compar-
Egg, hens 12.5 10.0 1.2 75.3 ison with episodic hunts for animal prey) of feeding oneself in
a hunting and gathering society. Gathering pulse seeds is par-
Based on data from FAOSTAT Database http://www.fao.org/ag/; Mohamed AA and
ticularly convenient compared with parts of other plants. Pods
Rayas-Duarte P (1995) Composition of Lupinus albus. Cereal Chemistry 72: 643647;
from grain legume plants are handy and easily grabbed com-
Kohajdova Z, Karovicova J, and Schmidt S (2011) Lupin composition and possible use
in bakery A review. Czech Journal of Food Sciences 29: 203211. pared with subterranean roots and tubers. No tools other than
fingers and palms are needed to grab and hang on to the large
seeds. Keeping both feet firmly on the ground is less daunting
(Tables 1 and 2). Pulses also contain high levels of dietary fiber than climbing trees for aerial fruits and nuts. The dense seeds
(
20 g fiber per 100 g compared with 12 g fiber per 100 g for are more filling than leaves and will satisfy hunger in the
wheat, 9 g fiber per 100 g for soybean, and 3 g fiber per 100 g for absence of meat if hunting is unsuccessful. As a bonus to
 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview 267

those concerned about times of scarcity, the seeds can be dried
and stored for a period of months or more without loss of food
value when eaten or viability when planted. Pulse seeds are
large relative to cereals, so they are convenient to handle for
food preparation and planting. Because of their convenience,
availability, and composition, wild pulses were an accessible,
nutritional contributor to early hunting and gathering societies
across the world.
The wild progenitors from which pulses were domesticated
have been identified, with the exception of Vicia faba. The
archeological record for some pulses, particularly Phaseolus
species and Lens culinaris, is more complete than for others
such as Vigna unguiculata and Cajanus cajan. In whatever man-
ner they were developed, the domesticates quickly spread
across regions and continents, particularly to regions lacking
a wide range of highly desirable, locally domesticated food
crops such as Europe and North America. Over time, domesti-
cation, selection, and trade resulted in a wide variety of pulses
grown, contributing protein to human diets across the globe.
Pulse and cereal grain domestication appear to have taken
place at roughly the same time in human history, which sug-
gests that these two foods may be ancient partners in providing
human nutrition. While cereals provided most of the energy
and much of the protein requirements of humans, pulses
played a strong supporting role as a contributor of concen-
trated dietary protein. Concurrent pulse and cereal production
and their dietary consumption resulted in more sustainable
farming systems, as well as better nutritional balance, than
would have been derived from production and consumption
of cereals alone.
Consumption
Current grain legume consumption in many parts of the Old
World and New World appears to be affected by cultural iner-
tia; past consumption patterns in those regions are echoed in
the present. While pulses are common components of diets
around the world, the extent to which they contribute to the
diet correlates closely with the historic availability and accep-
tance of animal protein. Ancient civilizations that consumed
more pulses were those where animal protein was less available
or acceptable. They included those that maintained high
human population densities and overwhelmed local supplies
of wild and domestic animal protein (East Asian Old World
civilizations), those where few animals were domesticated
(Aztec and Mayan New World civilizations), and those where
predominant religious practices excluded animal protein con-
sumption (Hindu veneration of the cow in South Central
Asia). Even today, those regions of the world consume more
pulses than elsewhere.
Besides influencing the amount of pulse contribution to the
diet, cultural inertia expresses a parallel effect on the pulse
species consumed within a region. Those pulses with long
records of use in a particular country or region of the world
continue as dietary preferences today (Table 3). The common
bean, Phaseolus vulgaris, appears to be the most universally
consumed pulse species.
Early in the Common Era, the 40-day fast of Jesus in the
desert became the standard reference for monks belonging to
the Roman Church for the custom of mortification of the body.
The fasting monks were required to remain engaged in con-
ducting the monasterys daily functions; thus, the elders of the
monastic community began, in the fourth century, to standard-
ize the fast diet in order to ensure that their workforce received
at least the nutrition required to keep the body and soul intact
but without arousing earthly sensuality in monks eating
flavorful food. A satisfyingly dull puree of pulses, accompanied
perhaps by a vegetable soup, became typical of the single meal
allowed during fasts. Monks certainly had their fill of pulses as
they contemplated eternity; the fifth-century monasteries
scheduled
230 fast days per year.
Before widespread adoption in seventeenth-century Europe
of cuisine innovations such as leavened bread, people of medi-
eval societies typically ate warm servings of porridge twice
daily. These plates or bowls of boiled gruel incorporated read-
ily available and affordable cereal grains such as barley and
wheat and also pulses such as the wild-growing or domesticated
lentils first found in the Eastern Mediterranean and West Asia.
Meat consumption declined with increasing population,
decline of wild game stocks, and movement of people into
towns and cities. World trade, natural sciences, and the influ-
ence of aristocratic culture began to influence diets in the late
1600s. Three-a-day meals became common across socio-
economic groups, and cookbooks spread food culture through-
out the urban and rural societies. New foods including spices,
potatoes, and a thickener made by browning flour in fat known
as roux made their appearance and elbowed the cereal-and-
pulse-based porridge out of popularity. The Protestant Refor-
mation deemphasized fasting, which further reduced the
consumption of sturdy but unglamorous pulse porridges.
Two vastly different problems influence contemporary
grain legume consumption: malnutrition in food-deficit

Table 3 Pulses consumed in various world regions

Country or region Pulse species commonly consumed

Mexico Phaseolus vulgaris, P. coccineus, P. lunatus, P. acutifolius, Vicia faba

North America Phaseolus vulgaris, P. lunatus
Brazil Phaseolus vulgaris, P. lunatus, Cicer arietinum, Vigna unguiculata
The Caribbean islands Phaseolus vulgaris, P. lunatus, Vigna unguiculata, Cajanus cajan
India Vigna radiate, V. mungo, Cajanus arietinum, Cajanus cajan, Pisum sativum, Lens culinaris, Lathyrus sativus
Africa Vigna unguiculata, Phaseolus vulgaris, P. lunatus, Vicia faba
The Mediterranean and Europe Vicia faba, Pisum sativum, Phaseolus vulgaris

Adapted from Smart J (1990) Grain Legumes: Evolution and Genetic Resources. Cambridge: Cambridge University Press.
268 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview

regions and diet-related diseases such as obesity, diabetes, and trend in developed countries between the 1960s and now is, if
heart disease in food surplus regions. In food-deficit regions, anything, to include slightly less pulse in diets. In contrast, the
increased consumption of pulse crops could provide a concen- increased protein and energy consumption between the 1960s
trated source of protein, vitamins, and micronutrients that and now reported in the least developed countries (as defined
complement traditional low-protein, high-carbohydrate grain by the Food and Agriculture Organization of the United
and root-based diets. Additionally, because of the legumes Nations) results from inclusion of both more animal protein
ability to support biological nitrogen fixation and its contribu- and more plant protein in the diets, including a small increase
tion to increased crop diversity, local farms could benefit from in pulse consumption. One aggravating reason for low pulse
agricultural systems that integrate pulse crops adapted to the consumption is flatulence resulting from intestinal fermenta-
regional environment. In food surplus regions, substitution of tion of complex sugars and dietary fiber. If gassiness causes the
plant-based diets for excessive meat consumption and con- consumer discomfort or embarrassment, pulse foods will be
sumption of foods rich in dietary fiber such as pulses are avoided when other choices are available.
among the strategies promoted to combat the dramatic As one might expect, food use of pulses is higher in least
increase in diet-related diseases. developed countries when compared with developed countries
(Table 6). However, the percentage of total domestic pulse
supply used for food in the least developed countries declined
Composition substantially over the past 40 years, while in developed
countries, it increased slightly. Feed use and food use are mutu-
Pulses, with roughly double the protein content of cereals, are ally exclusive, so in the least developed countries, there is a
damned with faint praise as the protein of the poor. They relatively low percentage of supply used for feed, while in devel-
contribute excellent nutrition to all, but because they are inex- oped countries, there is a high percentage used for feed. Table 6
pensive relative to meat, they are particularly beneficial to indicates that on average both the least developed and the devel-
subsistence farmers and those with low disposable income in oped countries exported more pulse seed than they imported,
food-deficient countries. In such circumstances, pulses can with the exception of developed countries in the 1960s. Trade
account for
11% of daily protein intake and 6% of the
daily energy in the human diet (Tables 4 and 5). The praise, Table 5 Energy per capita consumption in two decades by country
however, is faint and damning because this association with groupings (kcal per person per day)
poverty, a connection perhaps firmly established with the por-
ridges of the Middle Ages, stigmatizes pulse foods among those Least developed
with or aspiring to have higher income who increasingly favor countries Developed countries
meat protein. In developed nations, protein consumption has
Protein sources 196271 200211 196271 200211
increased from the 1960s to now and is associated with
increased consumption of animal products. Scarcely, 2% of Total plant 1818 (93)a 2055 (92) 2176 (71) 2339 (71)
dietary protein in developed nations, and 1% of energy, Pulses 79 (4) 94 (4) 34 (1) 28 (1)
comes from pulses. In more affluent settings, pulses remain Cereals 1248 (64) 1324 (59) 1030 (34) 954 (29)
an underused alternative to high-priced animal protein. Inter- Starchy roots 171 (9) 202 (9) 146 (5) 132 (4)
estingly, as noted in the preceding text, at least one segment of Oilseeds 43 (2) 50 (2) 19 (1) 30 (1)
affluent society, lifestyle- or health-conscious eaters such as Total animal 142 (7) 180 (8) 886 (29) 935 (29)
Meat 54 (3) 67 (3) 277 (9) 352 (11)
vegetarians or those seeking the nutritional benefits of high
Eggs 3 (0) 5 (0) 42 (1) 44 (1)
dietary fiber, still values these plants. Even so, the consumption Milk 53 (3) 70 (3) 296 (10) 313 (10)
Fish 13 (1) 19 (1) 35 (1) 48 (1)
Table 4 Protein per capita consumption in two decades by country Grand total 1961 2236 3061 3274
groupings (g protein per person per day)
a
kcal capita1 day1 (% of grand total).
Least developed nations Developed nations Based on data from FAOSTAT Database http://www.fao.org.

Protein sources 196271 200211 196271 200211

a
Table 6 Pulse domestic production and utilization in two decades by
Total plant 40.3 (80) 45.0 (78) 43.9 (48) 42.5 (42) country groupings (% of total domestic supply)
Pulses 5.2 (10) 6.1 (11) 2.2 (2) 1.9 (2)
Cereals 29.1 (58) 31.2 (54) 31.2 (34) 27.8 (28) Least developed
Starchy roots 1.8 (4) 2.5 (4) 3.5 (4) 3.1 (3) countries Developed countries
Oilseeds 1.6 (3) 2.2 (4) 0.9 (1) 1.3 (1)
Total animal 9.9 (20) 12.6 (22) 48.1 (52) 58.0 (58) Source 196271 200211 196271 200211
Meat 4.1 (8) 4.7 (8) 19.6 (21) 25.8 (26)
Eggs 0.3 (1) 0.4 (1) 3.4 (4) 3.5 (3) Domestic production 109 108 101 137
Milk 2.8 (6) 3.8 (7) 17.5 (19) 19.5 (19) Food use 81 66 31 34
Fish 2.1 (4) 3.0 (5) 5.4 (6) 6.9 (7) Feed use 1 16 57 54
Grand total 50.2 57.6 92.0 100.5 Imports 1 8 9 24
Exports 10 15 7 60
a
g capita1 day1 (% of grand total).
Based on data from FAOSTAT Database http://www.fao.org. Based on data from FAOSTAT Database http://www.fao.org.
 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview
movement of pulses, however, is much more complex than can
be summarized using these aggregates of countries grouped by
economic development. The reader is encouraged to examine
trade data at the regional and country levels in order to discern
how pulses are moving around the globe.
Protein Complementation
Understanding amino acid balance is key to recognizing the
special contribution made by grain legumes to protein comple-
mentation in cereal-based diets. Different sources of plant pro-
tein are made up of characteristic concentrations of
indispensable (essential) amino acids and therefore contribute
upon digestion a different balance of amino acids for human
nutrition. Amino acid balance in a particular protein source can
be scored against that of a reference protein, typically hens egg
(regarded as well balanced in relation to adult human metabolic
needs) or milk (Table 7). No plant-based protein mimics the
indispensable amino acid balance present in eggs, so it follows
that each type of plant protein must exhibit a characteristic set of
limiting amino acids. For example, whole wheat flour and white-
seeded common beans generally have lower proportions of the
indispensable amino acids than eggs or milk does. In particular,
the amino acid lysine limits efficient utilization of wheat protein
in human metabolism, while the indispensable sulfur-
containing amino acid methionine limits bean protein. The
indispensable amino acid balance of wheat and bean is charac-
teristic of other cereal grains and pulses. Cereal grains are typi-
cally deficient in lysine but high in methionine, while the
opposite is the case for pulses: high in lysine and low in methi-
onine. Cysteine, a dispensable sulfur-containing amino acid
synthesized from methionine and serine, is in many references
listed along with methionine as a limiting amino acid of pulses.
Plant protein complementation, accomplished by consum-
ing protein from more than one plant source, can provide
humans with a balanced diet of amino acids approximating
that found in hens egg. Grain legumes, with their high concen-
tration of lysine relative to lysine-deficient cereals, and cereals,
with their high concentration of methionine and cysteine rela-
tive to that found in pulses, combine as ideal dietary partners,
because the amino acids making up their proteins are very good
complements. For example, at 0.7 g kg1 body mass per day, an
85 kg adult would require
60 g of protein per day containing
Table 7 Indispensable amino acid requirements in dietary protein and the amino acid composition of selected foods (mg amino acid per g protein)
Amino acid Infant requirement Adult requirement
Histidine 23 17
Isoleucine 57 23
Leucine 101 52
Lysine 69 47
Methionine cysteine 38 23
Phenylalanine tyrosine 87 41
Threonine 47 24
Tryptophan 18 6 16
Valine 56 29
Based on data from Anonymous (2002) Dietary Reference Intakes for Energy, Carbohydrates, Fiber, Fat, Protein and Amino Acids (Macronutrients), Chapter 10, p. 71. Washington,
DC: National Academies Press (online version available on p. 535 at the URL http://www.nap.edu/books/0309085373/html/).
269
at least 2820 mg lysine. This requirement could be satisfied by
consumption of 288 g of wheat (Triticum aestivum) and 124 g of
bean (Phaseolus vulgaris) (Tables 2 and 7). To meet the lysine
requirement through consumption of wheat alone, the same
adult would need to eat 960 g wheat containing
108 g total
protein. The intentional combination of cereals and pulses at
meals offers inexpensive (in comparison with meat) plant-based
diets with balanced amino acid composition.
While the central theme of this overview is the enormous
contribution made by dry grain legume seeds, and in particular
pulse seeds, to the human diet, these crops have other food
uses. Some species are used as vegetables, as either green mature
or immature seeds or pods, as sprouted seeds, or even as stocks
for fermentation. As noted earlier, soybean is the base of many
nonfermented and fermented foods such as tofu and tempeh.
Fermented pulse seeds primarily involving black gram (Vigna
mungo), Bengal gram (Cicer arietinum), and green gram (Vigna
radiata) are components of diets particularly in India as idli,
dhokla, and khaman. Fermentation, sometimes in combina-
tion with cereal grains, causes desirable reduction in beany
flavor, aroma, and consistency and improves digestibility.
Nonfood uses include dry seeds used as animal feed and
forage from pulse vegetation, in aquaculture feeds, and even
as ornamentals. In a Southeast Asian setting, well-established
shrubs of pigeon pea, a short-term perennial grown on field
banks or as an intercrop, as well as dry season-grown horse
gram and cowpea, can be intermittently grazed or cut for hay.
N concentration of the hay varies between 2.5% and 4.5%,
but livestock may need a few days to be accustomed to the
feed. Dairy cattle are reported to similarly consume and uti-
lize faba bean silage and grasslegume silage. From a cool
season perspective, the seeds of soybean, pea, lentil, faba
bean, and chickpea can be used as a protein source in live-
stock feed. Oil and fiber contents of the seeds affect the
metabolizable energy (MEn) values of pulses when fed to
various types of livestock. For example, the relatively low-
oil, high-fiber pea has a low (2570 kcal kg1) MEn when
fed to poultry, while for swine, the pea MEn is much higher
(3740 kcal kg1). The pulse seed amino acid profile for
monogastric feeds, while good, is inferior to soybean. Pea
and faba bean protein is highly digested, but some livestock
feeding studies have shown an increased excretion of endog-
enous protein, which may be due to seed antinutrients. Dry
roasting the whole seeds has been shown to improve
Hens egg Cows milk Whole wheat flour White bean flour
24 28 22 28
63 60 40 42
88 98 63 76
70 79 26 72
56 34 35 19
98 96 81 77
49 45 27 39
14 11 10
72 67 43 46
270 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview
digestibility and protein utilization. Pulses have great poten-
tial for use in aquaculture diets, but in this application, they
are hampered by their carbohydrates. However, field peas
(Pisum sativum), faba beans (Vicia faba), chickpeas (Cicer
arietinum), and vetch (Vicia sativa) seeds can be used in
aquaculture feeds for juvenile silver perch following de-
hulling, which removes much of the undesirable carbohy-
drate fraction.
Antinutritional Factors
Low digestibility hampers full utilization of grain legume pro-
tein. In part, the problem may result from the more rapid
discharge, relative to other foods, of digesting pulses from the
intestinal tract and reduced protein hydrolysis by gut enzymes.
However, antinutritional factors, compounds in pulses that
interfere with their digestion and metabolism, also play a
major role in restricting dietary utilization in some pulse spe-
cies. These compounds generally include proteinaceous mole-
cules such as protease inhibitors (particularly trypsin
inhibitors) and lectins and also nonproteinaceous compounds
such as tannins.
Most of the wild relatives of pulses contain toxins and
antimetabolites. One of the key features of pulse domestication
from wild forms is human selection for lower levels of these
compounds. Traditional food preparation techniques used for
pulse seeds result in reduction or elimination of these meta-
bolic impediments. Some modern food pulses today, in par-
ticular the Phaseolus species, can still contain sufficiently high
levels of antinutritional factors in their dry seeds to cause
digestive difficulties if eaten without proper processing. For
this reason, pulse seeds are not eaten raw, but processed in a
way (moist heat >100  C, sprouting, and fermentation) that
inactivates the antinutritional factors.
Protease inhibitors, a major class of antinutritional factors
in pulses, often inhibit the digestive enzyme trypsin but may
act more broadly by inhibiting chymotrypsin and other serine
proteases. The role of these inhibitors in the plants that pro-
duce them is uncertain but may involve defense against disease
and insects. An alternative explanation is that these com-
pounds might simply serve as protein reserves high in sulfur-
containing amino acids such as cysteine and coincidentally
inhibit human digestion. The primary concern regarding raw
pulse consumption is with very young humans and other
young animals that are more susceptible to the effects of pro-
tease inhibitors than adults. In vitro studies suggest that prote-
ase inhibitors can impair nutrient utilization and reduce
growth rates in these groups. In vivo animal studies indicate
that interference with growth occurs primarily when the level
of dietary protein itself is limiting. Pancreas activity can be
affected when dietary protein digestion is inhibited and high
levels of undigested protein interfere with appropriate pancre-
atic regulatory control. Since pulses are used in some human
infant diets, cooking must be done with particular care to
ensure that protease inhibitors are inactivated.
Lectins, another major class of antinutritional factors in
pulses, are proteins that bind to carbohydrates or to molecules
containing carbohydrates. This binding capacity allows them
to agglutinate red blood cells (lectins are sometimes called
phytohemagglutinins) of different animal species depending
on the specific receptors on the cell membrane surface. This
agglutination specificity provides a method of rapid detection
and classification. Lectins differ in the severity of their impact.
Some (mainly found in Phaseolus vulgaris) are classified as
toxic, others are considered only growth inhibitory (Phaseolus
lunatus), and still others essentially nontoxic (Pisum sativum,
Lens culinaris, and Vicia faba). The role of lectins in the plant is
not well established, although they appear to help the plant
defend itself against specific bacteria, fungi, and insects. They
may also aid processing and movements of glycosylated stor-
age materials during seed maturation, assist in the establish-
ment of the symbiotic relationship of legumes and rhizobia,
and promote cell growth and division. Or, like protease inhib-
itors, lectins may simply serve as storage proteins.
The effect of ingested lectins on human and other animal
metabolisms varies according to the lectin type, the species
ingesting the protein, and the age, nutritional, and health
status of that human or animal. To affect human metabolism,
plant lectins first must bind to epithelial cells in the gut. This
binding requires the presence of particular carbohydrate
groups on the lumen of the gut epithelial cells. Dietary lectins
strongly resist proteolytic degradation in the gut. Toxic dietary
lectins can modify hormone balance and lipid and muscle
metabolism. These modifications deplete stores of lipid, glyco-
gen, and protein and lead to weight loss. Some nontoxic lectins
bind to the gut epithelium but have no deleterious effect on
metabolism, while others exhibit only limited binding capac-
ity. Deleterious lectin effects appear to be reversible.
Tannins can form strong cross-linked complexes with die-
tary proteins and enzymes. The antinutritional impact of tan-
nins on digestion and metabolism is not fully understood.
Protease inhibitors and lectins are heat labile and
rendered innocuous by usual methods of cooking. Moist heat
treatment such as the traditional soaking of beans, followed
by cooking in boiling water is generally considered to be
sufficient to inactivate protease inhibitors. Soaking may help
leach out some tannins, but heat treatment will not reduce the
impact of that remaining in the seeds.
Particular grain legume species accumulate additional clas-
ses of antinutrients. For example, Lupinus albus seeds may
contain quinolizidine alkaloids that give the seeds a bitter
flavor and, if not successfully removed prior to ingestion, can
result in poisoning. A. hypogaea cotyledons contain low-
molecular-weight glycoproteins to which some individuals
exhibit allergic reaction upon contact and ingestion. Neuro-
and osteolathyrism, the former a reversible paralysis and the
latter a skeletal abnormality, have been linked to excessive
consumption of grass pea (Lathyrus sativus) seeds. Broad bean
(Vicia faba) antinutrient levels are low and do not require moist
heating or fermentation to inactivate but can induce favism, a
hemolytic anemia, in individuals with congenital deficiency of
glucose-6-phosphate dehydrogenase. Pulses of Mediterranean
origin such as peas (Pisum sativum), chickpeas (Cicer arieti-
num), and lentils (Lens culinaris) generally have low toxic and
antinutrient levels. While protease inhibitors have been found
in cowpeas (Vigna unguiculata), they are apparently not present
in serious levels, nor have harmful quantities of cowpea lectins
been reported. Vigna mungo and Vigna radiata seeds likewise
contain little in the way of antinutritional factors.
 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview
Agronomy
The Leguminosae family is enormous, with an estimated
16 00019 000 species in
750 genera. If the raw number of
species were to determine the greatness of a family, Legumino-
sae would be eclipsed only by Orchidaceae and Compositae. If
instead the economic value of a family were considered,
legumes would be second only to Gramineae. Taxonomists
place pulses in the legume subfamily Papilionoideae, mostly
in tribe Phaseoleae, but a few in the tribes Vicieae, Cicereae,
and Genisteae.
Cropping systems develop in response to the demand for
particular crops, soil nutrient status, soil physical structure, and
biotic competition and are constrained by available resources
(including climate) and knowledge. Pulses in temperate crop-
ping systems are commonly planted in monocultures as part of
a rotational sequence with maize (Zea mays), small grain
cereals, or occasionally forages or periods of fallow in dryland
areas. Since in these cropping systems pulses are typically cash
crops and occasionally feed crops, their inclusion in the rota-
tion and total acreage from year to year depends on the status
of cash markets and economic benefits of growing that pulse
crop relative to other crops, rather than with maintaining a set
crop rotation, maintaining particular soil characteristics, or
addressing biotic competition. Higher-value crop alternatives
may be readily substituted for pulses in the rotation in some
climates. National farm strategies with strong economic
impact, such as government subsidies or deficiency payments,
can affect pulse production by inclusion or exclusion of partic-
ular pulse crops from the programs. For example, in the United
States, the Agricultural Act of 2014 identifies specialty crops,
including pulse crops, for indirect support such as assistance
with marketing, promotion and trade, crop insurance, and
research and research-based information extended to growers.
While not a big producer of any of these pulse crops, the
United States is an exporter, and additional production stimu-
lated by the Agricultural Act could affect prices in other pulse-
exporting or pulse-importing countries.
Leguminosae will collaborate with genus Rhizobium bacte-
ria in an ecologically unique symbiotic relationship. This sym-
biosis supports biological nitrogen fixation and frees legumes
from the need to access soil sources of fixed nitrogen. Rhizobia
invade the legume root hairs and reside in cortical root swell-
ings (nodules) where nitrogen from the atmosphere is fixed for
eventual transport within the plant and assimilation into pro-
tein. Even though a legume can support the symbiotic relation-
ship, it will not occur unless the rhizosphere contains
compatible rhizobia. Strains of rhizobia differ according to
the particular legume hosts that they will inoculate and are
classified according to this host range.
Cropping systems in developing countries typically rely on
intercropping where pulses are planted along with other
species, commonly cereals and vegetables. The intercropping
strategy can provide subsistence farmers with several essential
foods from one plot of land, produce higher protein yield per
hectare, combine complementary canopy structures (low-
stature pulses with high-stature sorghum (Sorghum bicolor)
and maize (Z. mays)), balance utilization of soil moisture
following rainy periods, or reduce crop competition by weeds
and insects. It is unlikely that a nonlegume companion crop of
271
similar growth period will benefit directly from the pulse crops
nitrogen fixation, but perennials, crops with longer growth
periods, or subsequent crops may benefit from the nutrients
released through the decomposition of pulse crop litter. The
agronomic impact of a legume crop on the intercrop will
depend on the crop species (and even the cultivar) in the
intercrop, the extent of its inclusion in the plot, tillage, and
soil type.
Pulses can assist in soil improvement, particularly when
used as green manure crops when the whole plant is plowed
down into the soil. Fixed nitrogen and in some cases phospho-
rus are made available to subsequent crops, and the added
organic matter contributes to improved soil structure. Also,
the deep tap roots of some crops such as pigeon pea reduce
soil erosion and mine deep soil for nutrients that are returned
to the surface through leaf drop.
In sustainable shifting cultivation systems of intercropping,
a few years of crop production are followed by a decade or
more of bush fallow where cropping is suspended and native
vegetation regrows. Cultivated pulses grown during the crop-
ping years provide concentrated protein for human diets using
nitrogen fixed from the atmosphere and help maintain the soil
nitrogen status by not depleting fixed soil nitrogen. During the
bush fallow stage, indigenous legumes, including trees, con-
tribute to the restoration of soil nitrogen. When the human
population density is low, relative to available land, and bush
fallow periods are sufficiently long to restore soil nutrients,
shifting cultivation can be sustained. With increased popula-
tion or decreased land base, the cropping period must be
extended, bush fallow period reduced, and as a result, soil
nutrient status and crop productivity decline.
Since in most cases pulse crops are not alternate hosts for
cereal crop diseases, their use in crop rotations will help reduce
cereal disease inocula. However, pulses are certainly not
immune to pests; they are attacked by many fungal, bacterial,
and viral diseases and a range of sucking, boring, and chewing
insects. Also, pulses are typically poor competitors against
weeds, except perhaps under stress conditions such as drought
conditions where annual weeds are also checked.
At harvest time, pulses often disappoint and frustrate
growers by failing to meet yield potential. Pulses are often
considered to be extremely sensitive to environmental stress,
relative to the more robust cereals, a sensitivity which may in
part be associated with maintaining the symbiotic relationship
with Rhizobium. The RhizobiumLeguminosae symbiosis
requires energy supplied by the host plant to support the
metabolic needs of the bacterial colonies in return for assimi-
lated nitrogen. In the case of the pulses, which are typically
grown as annual crops, the symbiotic relationship must be
reestablished with every new crop. Farmers watching the
color of the growing annual pulse crops in their fields can
clearly see the lag time between planting and the establishment
of functioning nodules. For example, a temperate common
bean crop planted in a soil with compatible rhizobia will tend
to have a deeper shade of green about a month after planting, as
fixed nitrogen becomes available to the plant. During the early
portion of the growing season, the host plant pays the costs of
establishment and only later receives the benefits. Plant-
generated energy used to support the symbiotic relationship is
not available to the plant for dry matter production, so some
272 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview
amount of yield penalty should be anticipated. Effective nodu-
lation and nitrogen fixation depend on favorable interactions
between the host and appropriate Rhizobium genotypes and the
environment. If these interactions falter and nitrogen fixation
lags behind demand, dry matter production will suffer.
Final Words
Declining per capita pulse consumption, particularly given the
nutritional value of pulses and their affordability, is a cause for
concern. Although some regions in developed countries now
benefit from new pulse export opportunities and are increasing
production, this expansion reflects changing markets and
increased population, not increased dietary utilization. Much
more global impact could be accomplished by addressing con-
straints to consumption, including the social stigma associated
with eating poor pulses compared with rich meat, flatulence,
yield variation, and agronomic constraints to product quality
and stable high yield. Further increasing pulse consumption in
developing and low-income countries may be impossible
given the strength of the human drive toward diets associated
with affluence. Perhaps, the greatest initial gains could occur
among consumers in developed countries currently with very
low pulse consumption. The keys to higher consumption in
those countries may involve increasing the availability of
value-added pulses such as those produced by certified organic
methods or food quality assurance programs and highlighting
their inclusion in national and regional cuisine initiatives. In
tandem, these efforts to add value and visibility could impart
some cache to pulses and improve their appeal.
Then said Daniel to Melzar, whom the prince of the eunuchs had set
over Daniel, Hananiah, Mishael, and Azariah, prove thy servants, I
beseech thee, ten days; and let them give us pulse to eat, and water to
drink. Then let our countenances be looked upon before thee, and
the countenance of the children that eat of the portion of the kings
meat: and as thou seest, deal with thy servants. So he consented to
them in this matter, and proved them ten days. And at the end of ten
days their countenances appeared fairer and fatter in flesh than all
the children which did eat the portion of the kings meat. Thus
Melzar took away the portion of their meat, and the wine that they
should drink; and gave them pulse.
Daniel 1:1116 (RSV)
Exercises for Revision
What seed constituents are similar between pulses and
oilseed legumes? What seed constituents differ? Which
grain legumes seed constituents are roughly intermediate
between pulses and oilseed legumes?
How do seed constituents of grain legumes compare with
those of cereal grains?
Did domestication of grain legumes generally occur in the
same region of the world or in vastly different regions?
Provide examples that support your answer.
Identify at least two reasons why pulse consumption tends
to be high in regions of the world where meat consumption
is low.
Which pulse crop is the most universally consumed?
A porridge made of grain legumes and cereal grains was a
common food in medieval societies. What events and
cultural shifts resulted in the decline of porridge in diets?
When did this transition take place?
Review the change in protein consumption from pulses and
from meat in the 1960s compared with recent years. How
do the trends differ between developed and developing
nations?
Identify examples of protein complementation in your diet.
Do any of your examples involve combinations of cereal
grains and grain legumes?
What are examples of nonfood uses of grain legumes?
Identify a reason for and a reason against using grain
legumes for animal feed.
Why are pulse seeds not eaten raw? What types of proces-
sing are necessary and how do they change the grain?
How would you integrate grain legumes into a cropping
system? What benefit would they provide to the system?
Describe the symbiosis that is key to the role of grain
legumes in the cropping system.
What constraints to consumption must be addressed before
grain legumes can play a larger role in global nutrition?
Exercises for Readers to Explore the Topic Further
Can you outline a climate change scenario that would
influence the production and consumption of grain
legumes?
Genetic transformation is not a common strategy for the
improvement of pulse crop cultivars. Why is this? What
factors might encourage or discourage the development of
GMO pulse cultivars?
What are the key challenges in producing pulse crops in
organic production systems? What are the benefits?
Few fast-food outlets include pulse-based foods on their
menus. Why is this? What types of meals made from pulses
might have promise as a fast and nutritious food?
What types of research have been conducted on the impact
of pulse-based foods on cancer and obesity? What have
been the outcomes of those research projects?
How can the social stigmas associated with pulse-based
foods (poverty and flatulence) be overcome?
How could the legumeRhizobium symbiosis be improved?
See also: The Basics: Taxonomic Classification of Grain Species;
The Oilseeds: Oilseeds: Overview; Soybean: Overview; Grains
Around the World: Grain Production and Consumption: Overview.
Further Reading
Adams MW and Pipoly III JJ (1980) Biological structure, classification and distribution
of economic legumes. In: Summerfield RJ and Bunting AH (eds.) Advances in
Legume Science, pp. 116. Kew, UK: Royal Botanic Gardens.
Allen ON and Allen EK (1981) The Leguminosae: A Source Book of Characteristics,
Uses and Nodulation. Madison, WI: University of Wisconsin Press.
 THE LEGUMES AND PSEUDOCEREALS | Grain Legumes and Their Dietary Impact: Overview
Booth MA, Allan GL, Frances J, and Parkinson S (2001) Replacement of fish meal in
diets for Australian silver perch, Bidyanus bidyanus: IV. Effects of dehulling and
protein concentration on digestibility of grain legumes. Aquaculture 196: 6785.
Dembinska M (1986) Fasting and working monks: Regulations of the fifth to eleventh
centuries. In: Fenton A and Kisban E (eds.) Food in Change, pp. 152160.
Edinburgh: John Donald Publishers.
Duke JA (1981) Handbook of Legumes of World Economic Importance. New York:
Plenum Press.
FAO (2002) Agriculture 21. Agriculture Department, Food and Agriculture Organization
of the United Nations. Rome, Italy: FAO Publishing and Multimedia Service (http://
www.fao.org/ag/).
Huisman J and van der Poel AFB (1994) Aspects of the nutritional quality and use of
cool season food legumes in animal feed. In: Muehlbauer FJ and Kaiser WJ (eds.)
Expanding the Production and Use of Cool Season Food Legumes, pp. 5376.
Dordrecht, The Netherlands: Kluwer Academic.
Jansman AJM, Hill GD, Huisman J, and van der Poel AFB (eds.) (1998) Recent
Advances of Research in Antinutritional Factors in Legume Seeds and Rapeseed.
Wageningen, The Netherlands: Wageningen Pers.
Labuza TP and Erdman Jr. JW (1984) Food Science and Nutritional Health: An
Introduction. St. Paul, MN: West Publishing.
Marquardt RR and Bell JM (1988) Future potential of pulses of use in animal feeds.
In: Summerfield RJ (ed.) World Crops: Cool Season Food Legumes, pp. 421444.
Dordrecht, The Netherlands: Kluwer Academic.
Reddy NR, Pierson MD, and Salunkhe DK (eds.) (1986) Legume-Based Fermented
Foods. Boca Raton, FL: CRC Press.
273
Reeds PJ and Beckett PR (1996) Protein and amino acids. In: Ziegler EE and Filer Jr. LJ
(eds.) Present Knowledge in Nutrition, 7th edn., pp. 6786. Washington, DC:
ILSI Press.
Sinha SK (1977) Food Legumes: Distribution, Adaptability and Biology of Yield. Rome,
Italy: Food and Agriculture Organization of the United Nations.
Smart J (1990) Grain Legumes: Evolution and Genetic Resources. Cambridge:
Cambridge University Press.
Summerfield RJ and Roberts EH (eds.) (1985) Grain Legume Crops. London: Collins
Professional and Technical Books.
t Mannetje L and Jones RM (eds.) (1992) Plant Resources of Southeast Asia. No 4.
Forages. Wageningen, The Netherlands: Pudoc Scientific Publishers.
Toussaint-Samat M (1992) History of Food. Cambridge, MA: Blackwell Publishers
(translated from the French by Bell A).
Relevant Websites
http://www.fao.org FAOSTAT Database, a comprehensive resource for agricultural
production data, including than on pulse crops.
http://www.icarda.org The website of the International Center for Agricultural Research
in the Dry Areas, based in Syria, has useful information on pulse agronomy and
utilization.
 Pseudocereals: Overview
RJ Fletcher, University of Queensland, Gatton, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 488493, 2004, Elsevier Ltd.
Topic Highlights
A pseudocereal is a plant grown to produce starchy grain
suitable for human food, excluding the cereals, legumes,
oilseeds and nuts.
The term pseudocereal combines cereal, referring to
grains of grass species, with the prefix pseudo- meaning
false or resembling.
The major pseudocereals are grain amaranth, quinoa and
buckwheat.
o Amaranth (Inca wheat); the seeds are used sprouted,
toasted, ground into flour, baked, cooked as porridge
and popped.
o Quinoa is rich in protein, fat and fiber, with a good
balance of essential amino acids.
o Buckwheat (unrelated to wheat) is milled into grits for
breakfast food or roasted to be boiled, steamed or baked.
Most pseudocereals have smaller seeds than the major
cereal grains.
Many pseudocereals can grow in poor soils and situations
not suitable for other grain species.
A large number of species are described that have the
potential to be useful as pseudocereals.
Learning Objectives
Understand the range of grains in the category of pseudo-
cereals, and the potential advantages they offer with respect
to growth situations, utilization and nutritional attributes.
Introduction
Most human foods come from plants. Roots, underground
storage or propagation organs, stems, petioles, shoots, flowers,
or fruits are eaten. Fruits in the form of grains or seeds are the
major form of human food.
Almost all of the grains eaten by humans are cereals. A
cereal grain contains an embryo and usually a quantity of
endosperm, enclosed within a seedcoat. Cereals are one type
of monocotyledonous plant (all of which present a single
shoot on germination); pseudocereals are noncereals, compris-
ing noncereal monocotyledonous plants as well as dicotyle-
donous plants; the latter have grains containing no endosperm
but an embryo enclosing perisperm.
The usefulness of any grain as human food (its functional-
ity) depends on the quantity and quality of protein, starch, and
lipid (fats and oils) present in the tissues of the grain (embryo,
endosperm and/or seedcoat).
Grain functionality depends principally on the genetic and
environmental influences on protein, starch, and lipid.
274 Encyclopedia of Food Grains, Second Edition
The proportion of embryo, endosperm, or perisperm
within any grain is primarily determined by a plants genetic
make-up. The eventual composition of the grain can be influ-
enced by the natural growing conditions and the manner in
which the crop and its product have been managed. Important
environmental factors not only include the growing conditions
but also the method of harvesting, the manner in which the
grain was stored, the nature of any processing carried out,
including the extraction method used, and the presence of
any antinutritional factors.
Grains used in the human diet come from surprisingly few
crops.
Such crops are collectively referred to as the cereals
(including crops such as wheat, rice, maize, barley, oats, rye,
etc.), pulses (including bean, chickpea, cowpea, pigeonpea,
lentil, lupin, etc.), oilseeds (including canola, sunflower, saf-
flower, sesame, coconut, cottonseed, flaxseed, mustard, etc.,
as well as some pulse crops, such as soybean and peanut) or
pseudocereals.
There can be some confusion with the use of such crop
groupings, since each grouping is not mutually exclusive; for
example, some pulse crops are used as oilseed crops and the
term pseudocereal is sometimes used to refer to crops more
commonly identified as oilseeds or legumes.
Definition
The American Heritage Dictionary of the English language
defines a pseudocereal as any of several plants, such as qui-
noa, that do not belong to the grass family but produce fruits
and seeds used as flour for bread and other staples.
The Oxford English Dictionary defines the term cereal as a
name given to those plants of the family Graminaceae [now
Poaceae] or grasses which are cultivated for their seed as
human food . . . sometimes extended to cultivated leguminous
plants. The term pseudo is defined as false, pretended, coun-
terfeit, spurious, sham, falsely so called or represented. When
combined with another element, such as the term cereal,
pseudo is defined as indicating close or deceptive resemblance
to the thing denoted by the second element, without real
identity or affinity with it.
The term pseudocereal is best used to refer to a plant which
is grown as a crop to produce starchy grain suitable for
human food, excluding plants already classified in a grouping
such as the cereals (those species from the grass family
Gramineae), legumes (those species from the legume families
Leguminosae-Caesalpinioideae, Leguminosae-Mimosoideae,
and Leguminosae-Papilionoideae), oilseeds (those species
described in terms of the commercially useful lipid or tricylgly-
cerol content), or nuts.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00039-5
 THE LEGUMES AND PSEUDOCEREALS | Pseudocereals: Overview
Current Role of Pseudocereals
The pseudocereals are relatively unimportant on a global scale
today. However, they have been significant contributors to the
human diet in certain defined regions in the past and could be
developed again as important new crops. For example, pseu-
docereals may play a role in human nutrition for those who
have allergies to traditional cereals; for the primary producer,
pseudocereals can play a role in cereal rotations, reducing the
buildup of grass weeds, pests, or diseases.
The three best-known pseudocereal crops are grain ama-
ranth (Amaranthus caudatus; A. cruentus; A. hypochondriacus;
family: Amaranthceae), quinoa (Chenopodium quinoa subsp.
quinoa; Chenopodiaceae), and buckwheat (Fagopyrum
esculentum; Polygonaceae). Each is briefly described below,
from the viewpoint of functionality. A range of potential
pseudocereals from other plant families is then presented,
with dicotyledonous families followed by monocotyledonous
families.
Weeds
Many of the species listed as potential pseudocereals below are
currently regarded, in some areas, as weeds; one of them,
Amaranthus retroflexus, could be regarded as the worlds most
cosmopolitan weed.
A weed is defined as a plant which is growing in the wrong
place. If some of the potential pseudocereals listed below can
be commercialized, they would no longer be regarded as
weeds.
Warning
The historic use of any obscure plant by a culture for medicinal
or similar personal use does not mean that the plant is safe for
use as a human food. Poisonous or antinutritional factors exist
in many plants and their historic use was often complemented
by elaborate preparation to reduce the effect of any poison. As
an example, Australian aborigines were able to consume cycad
seeds and yams but only after extensive preparation, which
reduced the toxicity of the material.
Regular use of a plant over a lifetime by a group of people
may also render a plant relatively safe for that group of people,
whereas a newcomer who has never eaten the food can suffer
serious effects on eating it for the first time.
Amaranthaceae
Amaranthus caudatus, A. cruentus, A. hypochondriacus
(grain amaranth)
Amaranthus caudatus (Inca wheat, love-lies-bleeding) is native
to the northern higher-altitude regions of Bolivia, Peru, and
Ecuador, whereas Amaranthus cruentus (Purple amaranth) and
Amaranthus hypochondriacus (Princes feather) are native to
Guatemala and Mexico. The species were grown by pre-
Colombian civilizations, Aztecs, and indigenous US tribes
but have mostly been replaced by cereal crops.
Depending on the species, grain amaranth seeds are
sprouted, parched, toasted, ground into flour, baked, mixed
275
with sugar to make confectionery items, rolled into balls,
cooked as porridge or popped.
Grain amaranth has enjoyed a resurgence in recent years,
especially through the health food market, since its crude
protein is high (1419%), with a high lysine (up to perhaps
6% of the protein) and tryptophan content. It appeals to the
modern consumer since these essential amino acids are low in
cereals.
Grain amaranth is being developed as an energy food to be
combined with traditional cereal grains in breakfast food,
bread, multigrain crackers, pastes, pancake mixes, or popped
as a snack food product. Popping can, however, affect its
nutritional quality.
The grain is easy to digest and heat processing improves its
digestibility.
Amaranth starch (up to 69% of the grain) is principally
amylopectin; granules are relatively small (13 mm) compared
with cereals (330 mm) and have a higher solubility and gela-
tinization temperature, rendering a distinctive gel. The seed can
comprise as much as 10% oil; this oil contains squalene, which
is used in cosmetics manufacturing. Antinutritional trypsin
inhibitors occur in concentrations up to twice that observed
in wheat.
Potential pseudocereals in the Amaranthaceae family
Amaranthus dubius (spleen amaranth, khada sag), Amar-
anthus frumentaceous (poong keeray, tola kura), Amaranthus
tricolor (tampala), Amaranthus graecizans (spreading pig-
weed), Amaranthus blitum (livid amaranth), Amaranthus
quitensis (yuyo colorado, sangorache), Amaranthus retro-
flexus (redroot, pigweed), Amaranthus spinosus (spiny ama-
ranth): the seeds are eaten raw in India and other places or
mixed with other grains or processed into flour or other
products.
Achyranthes aspera (devils horsewhip, prickly chaff-flower):
the seeds have been used in desert areas in India as a famine
food.
Celosia argentea var. argentea (red-fox), Celosia argentea var.
cristata (cockscomb): the seeds of these species have been
used as famine foods.
Chenopodiaceae
Chenopodium quinoa subsp. quinoa (quinoa)
Chenopodium quinoa subsp. quinoa is native to the border areas
between Peru and Bolivia. It held a special position in the
agricultural and ceremonial life of the Inca people and was
grown in Colombia, Peru, the southern areas of Bolivia, the
northern areas of Argentina, and the central areas of Chile.
Quinoa grain has a protein content ranging as high as
22.8%, a carbohydrate content up to 77.4%, a fat content up
to 9.5% (and relatively stable), and a fiber content up to 5.8%.
The grain is higher in protein, fat and fiber, and lower in
carbohydrate content than most comparable cereals, due prin-
cipally to the proportional size of the embryo within the grain
(up to 30% of the grains gross weight, compared with 1% for
most cereals).
The proteins in quinoa are principally albumins and glob-
ulins, which are at higher concentrations than is normally
276 THE LEGUMES AND PSEUDOCEREALS | Pseudocereals: Overview
found in the major cereals (wheat, rice, or maize). The amino
acid balance (higher in histidine, lysine, isoleucine, methio-
nine, and cysteine content) and mineral content (calcium,
magnesium, phosphorus, potassium, and iron) of quinoa
grain is superior to most cereals. However, there are saponins
in the seedcoat which have antinutritional properties.
Potential pseudocereals in the Chenopodiaceae family
Chenopodium album (fat hen, lambs quarters). The seed is
sprouted or ground into flour for breads, pancakes, muf-
fins, and biscuits in Russia and China; the grain is reported
to have 16.1% protein, 6.9% fat, 48.9% carbohydrate, and
5.8% ash.
Chenopodium ambrosioides (wormseed, Mexican tea). In the
Himalayas, the seed is ground into flour and mixed with
warm water or roasted or added to alcoholic beverages; care
needs to be taken with its preparation and use.
Chenopodium berlandieri subsp. nuttalliae (southern huauzon-
tle). The seed is ground to a meal and used for bread or
gruel (a light form of porridge produced by boiling meal in
water or milk).
Chenopodium murale (nettle-leaf goosefoot, sowbane). The seed
is popped or parched and ground to make gruel.
Chenopodium pallidicaule (canihua). The seed is toasted and
ground into a flour and eaten as a breakfast food, mixed
with wheat flour in baked products or drunk as a beverage.
Chenopodium sp. (California Yokuts lambs quarters). This
selection was used as a semi-domesticated grain crop by
Yokuts Indian villages, in the Central Valley, California.
Atriplex canescens (fourwing saltbush). The seed is ground
into meal and used as flour, or as a beverage.
Atriplex confertifolia (shadscale). The seed is ground into meal
and used for bread or mush (a heavier form of porridge than
gruel, produced by boiling meal in water or milk).
Atriplex hortensis (orache). Flour is prepared from the seeds
and used in soup and muffins; the seed is reportedly rich in
vitamin A.
Atriplex lentiformis (quail bush) and Atriplex polycarpa (all
scale).The seeds are eaten by the Native American Pima
group, located in Arizona.
Atriplex patula (halberd-leaved saltbush). The seed is ground
and mixed with corn and steamed as meat balls by the Zuni
Indians.
Atriplex prostrata (hastate saltplant). The seed is ground into
flour for use in baked products.
Haloxylon sadicornicum.The seeds are mixed with other
grains for bread making in India.
Suaeda corniculata and Suaeda heteroptera. The seeds are
eaten raw in Manchuria.
Tecticornia verrucosa. The seeds are ground to flour by
Australian aborigines and baked.
Polygonaceae
Fagopyrum esculentum (buckwheat)
Buckwheat originated from south central China.
This crop is harvested with its green pericarp intact; once the
pericarp (hull) has been removed, the groat can be coarsely
milled into grits for breakfast food or roasted and sold in whole
or a granulated form to be boiled, steamed, or baked. Buck-
wheat flour itself is used in soba-style Japanese noodles,
polenta (a form of porridge), desserts, ice cream cones, dietetic
foods, and canned meat or vegetable products and as a com-
ponent (3040%) of the flour used in pancakes, breads, pasta
products, cakes and dumplings, often combined with wheat
flour.
It is expected that buckwheat flour could be used more
readily in specialty breads, pasta, extruded snack food, and
ready-to-eat cereals.
The proteins in buckwheat grain are principally globulins
with a small proportion of prolamins; buckwheat protein has
twice the lysine content of wheat or white rice. The grains are
relatively high in potassium, magnesium, phosphate, iron, and
vitamins B1 and B2.
Potential pseudocereals in the Polygonaceae family
Fagopyrum cymosum (perennial buckwheat) or Fagopyrum
tataricum (Tartary buckwheat). In the Himalayas, the seeds
are eaten or ground into flour.
Calligonum polygonoides (phog, phogalli).The seeds are eaten
raw in India.
Polygonum aviculare (knotgrass, gooseweed). The seeds are
eaten or ground into flour for use in cookies or pancakes;
they are sometimes mixed with wheat; care needs to be
taken with their preparation and eating.
Polygonum glabrum (sauri arak, jioti, aatlaria). The seeds are
parched and made into a kind of Indian sattu (a tradi-
tional weaning food mix often eaten by adults, usually
based upon roasted Bengal gram, wheat, and sugar with
spices added and served in a number of ways, including as a
porridge).
Polygonum orientale (Princes feather).The seeds are eaten in
China.
Polygonum plebeium (chimtee sag, raniphul, machichi). The
seeds are crushed on stones, cooked, and eaten in the
form of a damper; the dry seed contains 17.4% protein,
50.4% carbohydrate, 3% fat, and 16.2% fiber.
Rumex acetosa (sorrel dock). The seeds are used as a base for
bread in Scandinavian countries; care needs to be taken
with their preparation and eating.
Rumex acetosella subsp. acetosella (sheep sorrel). The seeds are
ground into flour to make a flat bread called sygrasbrod.
Rumex crispus (curled dock). The seeds are ground into meal
or flour and used in pancakes; care needs to be taken with
preparation and eating.
Potential Pseudocereals in Other Dicot Families
Other dicotyledonous families with potential pseudocereal
species producing starchy grains include: Bixaceae, Cactaceae,
Cannabidaceae, Caryophyllaceae, Chloranthaceae, Cistaceae,
Portulacaceae, and Trapaceae. Species of potential interest are
named in the following.
There are many other minor dicotyledonous families with
plants producing starchy grains which will not be referred
to again; these include: Ancistrocladaceae, Basellaceae, Buddle-
jaceae, Desfontainiaceae, Dioncophyllaceae, Droseraceae, Ery-
throxylaceae, Frankeniaceae, Gelsemiaceae, Leitneriaceae,
 THE LEGUMES AND PSEUDOCEREALS | Pseudocereals: Overview
Lennoaceae, Molluginaceae, Nepenthaceae, Nyctaginaceae,
Orobanchaeae, Pentaphragmataceae, Phytolaccaceae, Retzia-
ceae, Sarcolaenaceae, Sargentodoxaceae, Saururaceae, Schisan-
draceae, Stegnospermataceae, and Tamaricaceae. Most of these
families have very limited numbers of species.
Bixaceae
Bixa orellana (annatto). The seeds are used as coloring and
flavoring for meat, poultry, or fish; the seedcoat contains the
carotenoid, bixin, which is used as a tasteless coloring for
cheese, butter, margarine and chocolate and for color in soap
and skin products.
Cactaceae
Carnegiea gigantea (giant cactus). The seeds are ground into
flour and used to make soup, paste, and other products by
the Native American Papago group.
Ferocactus wislizeni (fishhook cactus). The seeds are ground,
parched, and used for bread and gruel.
Opuntia clavata (ishikana). The seeds are roasted by Native
Americans of the Acoma and Laguna groups; however, the
mucilage in the fruit can render the food objectionable.
Opuntia phaeacantha (bastard fig). The seeds are dried,
parched, and ground into meal for gruel and cakes.
Opuntia soehrensii (ayrampo). A violet dye is extracted from
the seeds and used as a food coloring.
Pachycereus pecten-aboriginum (cardon hecho hecho). In Mex-
ico, the seeds are ground and made into cakes.
Pachycereus pringlei (cardon). The seeds are toasted, ground,
and fashioned into balls.
Stenocereus stellatus (xoconochtli, joconostle). The seeds have
been eaten.
Cannabidaceae
Cannabis sativa (hemp). The seeds are parched, fried as cakes,
used in asanomi (in Japan), in the spice mix shichimi, in ale or
are eaten as sprouts.
Caryophyllaceae
Lychnia segetum. The seeds have been eaten as a famine food
in France.
Spergularia arvensis (corn spurrey). The seeds are used for
bread flour in Norway and Sweden.
Stellaria media (chickweed). The seeds are used as bread or as
a soup thickening in India.
Chloranthaceae
Sarcandra glabra (tea scent). The seeds are roasted and used as a
sesame substitute.
Cistaceae
Cistus ladanifer (labdanum). The seeds are ground and used for
cakes and breads.
277
Portulacaceae
Portulaca oleracea (purslane).The seeds are sprouted or ground
for use in gruel, cake, bread, or pancakes; the content of glycine
and tyrosine in the seeds is reportedly high; care needs to be
taken with their preparation and eating.
Trapaceae
Trapa natans (water caltrop, water truffle).The seeds are eaten
raw, roasted, boiled, fried, or ground into flour for bread
and sweet puddings.
Trapa bispinosa. The seeds are cooked and eaten in India.
Potential Pseudocereals in Monocot Families
Monocotyledonous families with potential pseudocereal spe-
cies include: Araceae, Cannaceae, Commelinaceae, Cyperaceae,
Marantaceae, Musaceae, Palmae, Pandanaceae, Pontederiaceae,
Typhaceae, and Zingiberaceae. Species of potential interest are
named in the following.
There are also many minor monocotyledonous families
with plants that produce starchy grains which will not be
referred to again; these include: Alismataceae, Anarthriaceae,
Aponogetonaceae, Bromeliaceae, Burmanniaceae, Butoma-
ceae, Centrolepidaceae, Costaceae, Cyanastraceae, Cyclantha-
ceae, Eriocaulaceae, Flagellariaceae, Haemodoraceae,
Hydatellaceae, Hydrocharitaceae, Joinvilleaceae, Juncaceae,
Juncaginaceae, Lemnaceae, Mayacaceae, Melanthiaceae, Naja-
daceae, Philydraceae, Potamogetonaceae, Rapateaceae, Rhipo-
gonaceae, Scheuchzeriaceae, Sparganiaceae, Triuridaceae, and
Velloziaceae.
Araceae
Peltandra virginica (arrow arum, tuckahoe). The seed produces a
bread similar to corn bread, with a cocoa flavor.
Cannaceae
Canna edulis (Queensland arrowroot). Immature seeds are
cooked in tortillas or cakes; the plant is used as a source of
arrowroot in Colombia.
Commelinaceae
Commelina benghalensis (keng, bokna, mu-Kengeria), Comme-
lina forskailii (kansura, kanshura). The seeds are ground into
flour for bread.
Commelina communis (spider wort, day flower), Commelina
obliqua (kena, keni, kana, kanjura). The seeds are eaten.
Cyanotis axilaris (vichaka, narido, damro, soltra). The seeds
are ground into flour for bread or bhadku, a mixture of
flour and salt.
278 THE LEGUMES AND PSEUDOCEREALS | Pseudocereals: Overview

Cyperaceae Elettaria cardamomum (cardamom). The seeds are used as

Cyperus bulbosus (theg, motha, mothabasa). The seed is
ground into flour and made into bread or ghes or rab
in India; Australian aborigines ate the seed raw or roasted.
Mariscus sieberianus (tall sedge). The seeds are made into
flour in China.
Scirpus lacustris (great bulrush).The seeds are ground and
mixed with meal for bread, mush, or pancakes.
Scirpus maritimus (chid, dila). The seeds are eaten raw or
pounded, made into flour and mixed with millet in India.
Scirpus validus (tall bulrush).The seeds are eaten.
Marantaceae
Thaumatococcus danielli (miracle berry). The seeds are used as a
sweetener but the extract does not withstand heat.
Musaceae
Ensete ventricosum (Abyssinian banana). The endosperm of the
seed is eaten in Africa.
flavoring.
Etlingera elatior (Phillipine wax-flower, ondje). The seeds are
eaten raw in Malaysia.
Conclusion
In the past, the pseudocereals have represented an important
component of the diet of many people in a wide range of
cultures. The representative list of potential pseudocereals
included above demonstrates the opportunities for future
development of functional foods that exist amongst the pseu-
docereals, based on the usefulness of these foods in the past. In
addition, the reputation of so many of the species as weeds
indicates that they will probably have growth characteristics
which should assist with their development into modern com-
mercial production systems.

Exercises for Revision

Palmae/Arecaceae
Hyphaene thebaica (gingerbread palm). The kernels are crushed
Pseudocereals constitute a diverse group. How is this group
defined?
and mixed with porridge; the edible portion of the kernel has
2.8% protein, 74.7% carbohydrate, 0.4% fat, and 11% crude
What are the most valuable attributes of the pseudocereals?

fiber.
Contrast and compare the characteristics (grain yield and
utilization value) of the three main pseudocereals: ama-
ranth, quinoa and buckwheat.
Pandanaceae Consider a further six potential pseudocereal species (e.g.,
chia) and indicate what beneficial attributes they offer.
Pandanus tectorius (Nicobar breadfruit, screwpine). The seeds are
edible but only after careful preparation.

Pontederiaceae
Exercises for Readers to Explore the Topic Further

Pontederia cordata var. cordata (lance-leaf pickerel weed). The Find out the potential situations in which some of the less
seeds are eaten raw, boiled or parched, and ground into flour common pseudocereals might become useful, considering
for bread. growth vigor and seed usefulness (including anti-
nutritional properties, such as saponins).

Typhaceae
Amaranthus retroflexus is considered a weed in some situa-
tions, but it appears to have some potential for animal feed,
Typha angustifolia (narrow-leaf cattail). The seeds are roasted possibly even for human food. Investigate the situations in
and eaten. which its potential value might be harnessed for good.

Zingiberaceae
See also: Food Grains and the Consumer: Grains and Health;
Aframomum angustifolium (Madagascar cardamom). The
Food Grains and Well-being: Functional Foods: Dietary
dried seeds are used like pepper or added to coffee.
Fibers, Prebiotics, Probiotics, and Synbiotics; Food Grains:
Aframomum melegueta (Melegueta pepper). The seeds are used
Intolerance, Allergy and Diseases: The Gluten-Free Diet;
as flavoring for beverages, meats, ice cream, candy, and
Grain Composition and Analysis: The Composition of Food
bread.
Grains and Grain-Based Products; Non-food Products from
Amomum compactum (round cardamom). The seeds are used
Grains: Cereal Grains as Animal Feed; The Basics: Grain and
in cakes in Malaysia.
Plant Morphology of Cereals and how characters can be used to
Amomum subulatum (Indian cardamom). The seeds are used
identify varieties; Taxonomic Classification of Grain Species; The
for flavoring in char masala, in Afghanistan.
Grain Crops: An Overview; The Legumes and Pseudocereals:
Amomum xanthioides (bastard cardamom). The seeds are used
Amaranth: Overview; Buckwheat: Overview; Quinoa: Overview.
as flavoring in liqueurs in China.
 THE LEGUMES AND PSEUDOCEREALS | Pseudocereals: Overview
Further Reading
Belton PS and Taylor JRN (2002) Pseudocereals and Less Common Cereals. Berlin:
Springer.
Crawford M (2000) Edible Plants for Temperate Climates. Totnes, Devon: Agroforestry
Research Trust.
Facciola S (1998) Cornucopia II. A Source Book of Edible Plants. Vistas, CA: Kampong
Publications.
Hegarty MP, Hegarty EE, and Wills RBH (2001) Food Safety of Australian Plant
Bushfoods. Canberra, ACT: Rural Industries Research and Development
Corporation.
Mabberley DJ (1997) The Plant-Book. A Portable Dictionary of the Vascular Plants, 2nd
edn Cambridge: Cambridge University Press.
279
Wiersema JH and Leon B (1999) World Economic Plants. Boca Raton, Florida: CRC
Press.
Williams JT (1995) Cereals and Pseudocereals. London: Chapman and Hall.
Relevant Websites
Biodiversity Research Collection, University of Connecticut - The families of flowering
plants: descriptions. http://www.biodiversity.uconn.edu.
http://www.pfaf.org/ Plants for a future.
hort.purdue.edu http://www.hort.purdue.edu Center for new crops and plant products,
Purdue University and Famine Foods database.
 Lupin: Overview
DS Petterson, Department of Agriculture, Nedlands, WA, Australia
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp 166174, 2004, Elsevier Ltd.
Topic Highlights
Origin of lupins
Grain morphology and composition
Lupin processing
Food uses of lupins
Allergenicity
Feed uses
Learning Objectives
To understand the structural and compositional aspects of
lupins
To understand the origin and uses of various types of
lupins
To appreciate the requirements for and types of lupin
processing
To understand the various antinutritional factors present in
some lupins
Definition and Origins of Lupins
The seed or grains of domesticated Lupinus species are generi-
cally called lupins. The same term is used widely in the com-
munity in referring to wild and domesticated plants and the
seed/grain of all Lupinus species. The genus Lupinus belongs to
the Fabaceae, a family of legumes. There are 12 lupin species,
all large-seeded, native to Europe and the Mediterranean
regions. Three of these are now fully domesticated. Lupinus
albus, the European white lupin or albus lupin has a white
flower. Wild types of L. angustifolius, the narrow-leafed lupin,
have a blue flower: however, modern cultivars bred for low
alkaloid content in Australia have a white flower and are
known widely as the Australian sweet lupin (ASL). L. luteus,
the yellow lupin (YL), has a yellow flower. Their grain is widely
used in the animal feeds industry and to a lesser extent as a
food (or food ingredient). There are
100 lupin species native
to the Americas. They are mostly small-seeded, and only L.
mutabilis, the pearl or Andean lupin, has been used as a food
source. Worldwide, several species are being domesticated for
possible use in agriculture. The main value of lupin crops to
farmers is their replenishment of soil nitrogen, providing a
disease break for cereals in crop rotation systems and a mar-
ketable grain. Lupins tend to grow relatively better on poorer
soils than most other crop species.
Lupins are also well known for their spectacular flowers.
The Russell lupin, a hybrid derived from L. polyphyllus, is widely
used as an ornamental flower, particularly in Europe. Parts of
the southern highlands in New Zealand are a blaze of color
from the alpine lupin, a variant of the Russell lupin, in
280 Encyclopedia of Food Grains, Second Edition
springtime and early summer. The (bluebonnet) flower of
L. texensis is the floral emblem of Texas.
It is widely accepted that Lupinus evolved from the tropical
and subtropical Sophoreae, a primitive tribe of the subfamily
Papilionoideae (pea-flowering plants), 40 Ma. Nowadays, wild
lupin species cover almost all climatic zones: sub-Arctic Alaska
and Iceland, Mediterranean and semidesert regions, the high-
lands of East Africa, Mexico and the Andes, and the subtropical
lowlands of the eastern Americas.
Grain Morphology and Composition
Lupins have a typical dicotyledon structure. Their thick seed
coat (hull or testa) composes
30% of the seed weight for L.
luteus, 25% for L. angustifolius, 15% for L. albus, and 12% for L.
mutabilis. This is considerably higher than for most domesti-
cated grain species. Within the cotyledons (kernels), energy is
mostly stored in the form of thickened cell wall material
(
25% of the cotyledons) and oil bodies (
614%). There
is virtually no starch in any of the lupin species. This is in
marked contrast to crops such as field peas and chickpeas,
which can have 5070% of the cotyledon weight as starch
and have low protein and oil content, and the soybean with
1520% oil, some starch, and a high protein content. Their
crude protein content ranges from
28% to 42%. Proximate
analyses for the whole grain of the major domesticated species,
and the Andean lupin, are shown in Table 1. There are some
wild and partly domesticated Lupinus lines containing up to
45% crude protein and up to 21% oil; others have as little as
20% crude protein and 3% oil.
The thick seed coat, which is mostly cellulose and hemicel-
lulose, of Lupinus species means that it is important to also
consider the composition and nutritional value of their coty-
ledons: these data are in Table 2.
The major lupin proteins are a group of globulins, called
conglutins. The three major ones are conglutin a (which sed-
iments at
1112S and is analogous to the legumin of peas
and glycinin of soybeans), conglutin b (which sediments at

7S and is analogous to the vicilin of peas and conglycinins a
and b of soybeans), and conglutin g (a sulfur-rich protein that
sediments at
2S and is analogous to the conglycinin of
soybeans). They compose
85% of the total protein and
have similar sizes and physical properties to the storage pro-
teins of other grain legume species. The remaining 15% of
proteins are albumins, which are soluble at pH 5 and vary in
size from
6000 to 117 000 Da. There is evidence for the 2S
globulin actually being an albumin: depending on the species,
this fraction consists of four (L. luteus), five (L. albus), or six
(L. angustifolius) proteins. For more details on lupin and other
legume proteins, see CEREALS | Protein Chemistry.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00036-X
 THE LEGUMES AND PSEUDOCEREALS | Lupin: Overview 281

Table 1 Chemical composition of the major lupin species (g kg1 as Table 3 Essential amino acid profile for major lupin species
received)a (% in grain)a

L. angustifolius Amino acid L. albus L. angustifolius L. luteus L. mutabilis

(Australian sweet L. mutabilis
L. albus lupin, L. luteus (pearl lupin, Arginine 4.7 3.6 4.4 4.6
(albus lupin, narrow-leafed (yellow Andean Cystine 0.5 0.5 0.7 0.7
white lupin) lupin) lupin) lupin) Histidine 0.6 0.8 1.4 1.3
Isoleucine 1.4 1.2 1.5 2.0
Moisture 86 85 94 71 Leucine 2.3 2.1 2.8 2.9
Protein 361 322 414 460 Lysine 1.6 1.4 2.1 2.5
(N  6.25) Methionine 0.2 0.2 0.4 0.3
Ash 33 28 37 40 Phenylalanine 1.2 1.1 1.8 1.7
Crude fat 91 58 57 110 Threonine 1.2 1.0 1.5 1.8
Crude fiber 104 150 127 110 Tryptophan 0.4 0.3 0.8 0.3
ADF 152 197 195 Tyrosine 1.7 1.1 1.2 1.4
NDF 171 227 230 Valine 1.4 1.2 1.3 1.7
TDF 390 284 320
a
Lignin 15 7 5 Data are averaged from values obtained worldwide.
Starchb <10 <10 <10
Gross energy 19.0 18.6 18.6
(MJ)
Table 4 Nonstarch polysaccharide components for major lupin
ADF, acid detergent fiber; NDF, neutral detergent fiber; TDF, total dietary fiber.
a species (g kg1 in grain)a
Average of worldwide data.
b
Most analysts report no starch present. L. albus L. angustifolius L. luteus
(albus lupin, (Australian sweet lupin, (yellow
Table 2 Chemical composition of the kernels of major lupin species white lupin) narrow-leafed lupin) lupin)
(g kg1 as received)a
Rhamnose 13 10 10
L. albus L. angustifolius L. luteus Arabinose 67 50 70
Xylose 22 10 21
Moisture 75 100 85 Mannose 1 1 1
Protein (N  6.25) 401 400 530 Galactose 202 170 133
Ash 36 27 45 Glucose 23 13 42
Crude fat 114 66 73 Uronic 50 29 43
Crude fiber 150 88 195 acids
ADF 62 71 220 NSP 390 284 320
NDF 140 77 300 (total)
TDF 440 380 420
a
Lignin 30 7 Data are averaged from values obtained worldwide.
Gross energy (MJ) 20.4 18.9 19.7

ADF, acid detergent fiber; NDF, neutral detergent fiber; TDF, total dietary fiber.
a water, about eightfold by weight for lupins, and maintain
Average of worldwide data.
Typical amino acid profiles for the domesticated lupin spe-
cies, compared with the FAO standards for infants and others,
are shown in Table 3. The deficiencies in lysine and methio-
nine are common to most grain legumes.
The seed hulls and cotyledons contain different types of
carbohydrate. Lupin hulls are predominantly composed of
structural polysaccharides: cellulose, hemicelluloses, and pec-
tins. The main carbohydrate reserves of the cotyledons are the
nonstructural polysaccharides of the cell walls, with the main
components being galactose, arabinose, and uronic acid. These
complex compounds are referred to as nonstarch polysaccha-
rides (NSPs). Their structure, based on a rhamnogalacturonans
backbone, is not yet fully defined. The water-soluble portion,

5% of the seed, is considered to have an antinutritional effect
due to its viscous nature and effect on intestinal transit time.
The insoluble NSP,
30% of the seed, has a minimal effect on
nutrient utilization by monogastric species. An important attri-
bute of insoluble NSP is their ability to hold large quantities of
normal gut motility. The profiles for various species are
shown in Table 4. The oil content of lupins varies from

15% in the pearl lupin down to 4% in the YL. There are
differences in individual fatty acid profiles; they are typically
high in oleic and/or linoleic acids and contain a total of
75%
unsaturated fatty acids (cf.
85% in olive oil). The oil is heat-
stable and has a high antioxidant capacity.
Variation in the content of major essential minerals in
lupins, up to
30% of the mean, is typical of legume species,
probably reflecting both genotype and environmental factors
(Table 5). The essential trace mineral content of lupin species
not only is influenced by genotype but also tends to reflect the
soil types on which they are grown. For example, the selenium
content of lupin seeds grown in Western Australia varies
according to species and rainfall zone. Where the species
were grown together, the selenium content of L. albus > L.
luteus > L. angustifolius. In all cases, the higher the rainfall, the
lower the selenium content. Similar variations apply to the
content of copper, cobalt, and zinc in lupins. The accumula-
tion of manganese by L. albus is well documented with most of
282 THE LEGUMES AND PSEUDOCEREALS | Lupin: Overview

Table 5 Mineral content of major lupin species (range of data sets) Table 6 Antinutritional factors in the major lupin speciesa

L. albus L. angustifolius L. luteus Botanical name L. albus L. angustifolius L. luteus

1 1
Mineral content (g kg ) Total alkaloids (mg kg ) <200 <200 200500
Calcium 1.23.3 1.53.1 1.83.0 Oligosaccharides (%) 7.5 5.2 12.3
Magnesium 0.91.6 1.12.0 2.23.2 Saponins (mg kg1) <1 570 55
Phosphorus 2.59.0 2.14.3 3.47.3 Condensed tannins (%) 0.01 <0.01 0.02
Potassium 2.811.1 6.6109.1 8.811.6 Lectins (activityb) Trace Trace Trace
Sulfur 2.12.7 1.52.9 4.04.9 Trypsin inhibitors (mg g1) 0.13 0.14 0.29
Sodium <0.11.1 0.31.1 <0.1 Phytate (%) 0.79 0.58 0.96
Mineral content (mg kg1)
a
Copper 3.18.1 2.56.8 5.912.0 Data are averaged across worldwide figures.
b
Iron 2144 31150 5287 Lectin activity is usually measured as units of agglutinating activity against red cell
Manganese 233772 6.776 2559 preparations from the blood of various mammalian species. Several authors argue that
Molybdenum 0.83.1 0.72.9 there is no significant activity in Lupinus species; some have detected activity against
Zinc 2238 2445 3982 specially sensitized cells or against species not considered significant for human
Mineral content (mg kg1) concern.
Cobalt 10430 10260 n.a.
Selenium 20360 18240 n.a.
effect of phytate in the diet depends on the overall composi-
n.a., not available.
tion of the food particularly the amount and types of protein
content and the total mineral content. Human studies have
the low values coming from crops grown on sandy soils and shown similar rates of absorption of phosphorus from lupin-
high values coming from crops grown on heavy red clays. In based foods and comparable soy products. Germination
contrast, the manganese content of L. angustifolius is low, from lowers the phytate content by
60% and fermentation
9% to 30% the amount in L. albus grown at the same site by
80%.
(Table 5). Tannins are compounds of plant origin with molecular
A survey of the cadmium and lead contents of L. angustifo- weights ranging from 500 to 2000 Da and with one to two
lius (178 samples) and L. albus (12 samples) grown in Australia phenolic hydroxyl groups per 100 Da. This enables them to
found that no samples exceeded the Codex Alimentarius limits form cross-linkages between proteins and other macromole-
of 0.1 and 2.0 mg kg1, respectively. There can be some accu- cules. There are two subgroups of tannins. Hydrolyzable tan-
mulation of cadmium by L. luteus, which may be a conse- nins typically have a central glucose core with the hydroxyl
quence of the different root architecture in this species and groups being wholly or partly esterified with gallic acid or
the soils in which it is grown. hexahydrodiphenic acid. Condensed, nonhydrolyzable tan-
Legume grains contain a range of compounds, traditionally nins are higher oligomers of flavan-3-ols with varying degrees
known as antinutritional factors (ANFs) or antinutrients, with of substitution. Their astringent taste and ability to precipitate
apparent untoward effects on species ingesting them. These are proteins, resulting especially in the inactivation of gut
also known as bioactive compounds. These compounds had a enzymes, give the tannins their antinutritional role. In lupins,
protective role in evolution by protecting against predators and as with other grain legumes, the tannins are concentrated in the
may have other important functional roles. Plant breeders have seed coats (hulls), and the simple act of dehulling will mini-
mostly reduced their content to a point of balance between mize any adverse effects. The concentration of condensed tan-
lowering the content of ANFs, to enhance the nutritional value nins, those most responsible for protein binding, is so low in
of the grain, and retaining the defensive or, otherwise, func- lupins that it is unlikely to affect human or animal nutrition.
tional role. While some ANFs have a negative effect on feed Saponins are plant glycosides in which the nonsugar moi-
utilization by farmed livestock, they can also have beneficial ety is a steroid or a triterpenoid compound. Their bitter taste
effects in humans, such as preventing the development of some acts as a feed deterrent, and they have a secondary antinutri-
forms of cancer and of osteoporosis. A summary of data for the tional effect by increasing the permeability of small intestinal
ANF content of lupins is shown in Table 6. mucosal cells. This causes a loss of essential electrochemical
The content of proteinaceous ANF in domesticated lupin concentration gradients, facilitating the uptake of materials to
species is very low. Typically, trypsin inhibitor activity is which the gut would not normally be permeable. Only traces
< 0.3 mg kg1, and chymotrypsin inhibitor activity, of saponins are present in L. albus, while concentrations in L.
<0.6 mgkg1. Lectin activity is virtually nonexistent in all angustifolius and L. luteus range from 55 to 730 mgkg1. Sapo-
lupin species. So, unlike most legume grains, lupins do not nins are generally harmless to humans: some are claimed to be
need to be heated to denature the proteinaceous ANF and beneficial in lowering blood cholesterol levels and protecting
make them safe for consumption by humans or animals. against coronary heart disease. The concentrations in lupins are
Phytate (inositol hexaphosphate and lower substituted lower than in many other legume species.
homologues and their salts) contributes about one-half of the Lupin oligosaccharides are higher a-galactosides of sucrose.
total phosphorus content in lupins. This is rapidly mobilized Raffinose has one galactose moiety linked to sucrose through
upon germination. Phytate can form insoluble complexes with an a-1,4 bond, while stachyose has two, verbascose three, and
divalent cations, particularly Ca2 and Zn2, thus making ajugose four. These compounds cannot be metabolized by
them less available for absorption and utilization. The net humans and other nonruminant species, and they undergo

bacterial fermentation in the colon to produce carbon dioxide,
methane, and hydrogen. This causes abdominal discomfort
and cramps and leads to flatulence, which seems to be a
major reason for the low interest in consuming grain legumes
in many societies. The oligosaccharides are a rich source of
nutrients for bifidobacteria in the colon. This can have a ben-
eficial effect, as the bifidobacteria counter the activity of putre-
fying bacteria and reduce their production of harmful, and
possibly carcinogenic, fermentation products. One further
source of confusion about the significance of oligosaccharides
is the inclusion by some authors of sucrose in their total for
oligosaccharides. This is nutritionally confusing as sucrose can
be absorbed from the stomach and upper digestive tract of
humans and nonruminant animals.
Isoflavones have generally been regarded as ANFs because
of their negative effects on fertility in ruminants grazing pas-
ture medics and subterranean clovers. However, they are now
recognized as preventative agents against some forms of cancer
and osteoporosis in women. Isoflavones are present in green-
leaf material and in the hypocotyls of germinating seeds
(sprouts). There is conflicting evidence as to their presence in
mature grain.
The lupin alkaloids are usually bicyclic, tricyclic, or tetra-
cyclic derivatives of quinolizidine (Figure 1). The individual
alkaloids and their concentrations vary widely between,
and sometimes within, species (Table 7). Grain of modern
domesticated lupin cultivars typically contains less than
200 mg kg1 alkaloids. In contrast, grain from bitter wild
types that still exist in many countries may contain from
5000 to 40 000 mg kg1 alkaloids. There have been human
deaths associated with the consumption of grain from bitter
wild types of lupin. One would need to eat
10 kg of the
modern varieties of lupin grain in a very short time to absorb
sufficient alkaloids to be at risk. The quinolizidine alkaloids
have a short half-life in humans and are excreted largely
unchanged in the urine of >90% of humans. Traces of
N the cotyledons to make flour or to separate it into protein-rich
Quinolizidine
Figure 1 Structure of quinolizidine.
Table 7 Quinolizidine alkaloids in the major lupin species (% in
alkaloid fraction)a
L. albus L. angustifolius L. luteus L. mutabilis
Albine 15
Lupanine 70 70
Multiflorine 3 meal or fine flour for nonruminant feeds. The particle size
13-Hydroxylupanine 8 12
4-Hydroxylupanine 1 7
Angustifoline 10
Lupinine 60
Sparteine 30
a
Data are for modern cultivars or developing lines of these species. Up to 10% gramine,
an indole alkaloid, is present in L. luteus.
THE LEGUMES AND PSEUDOCEREALS | Lupin: Overview 283
gramine, an indole alkaloid, occur in some lines of L. luteus,
but the significance of these levels is not known.
The only mycotoxin associated with lupins to date is the
family of phomopsins, a group of linear hexapeptides with an
antimicrotubule effect, which is most pronounced in the liver
and kidneys. They are capable of disrupting mitosis and of
reducing the activity of motile cells and organisms, rendering
their host prone to secondary infections. There is no relation-
ship between lupinosis and lupin alkaloid poisoning that can
occur when sheep graze bitter lupins and their stubble or graze
the perennial lupins of North America. Grains that contain
phomopsins are smaller and less dense than normal-colored,
nontoxin-containing grain and can be easily removed from the
rest of the harvested grain by grading, using screens, gravity
tables, and aspirators. A rapid enzyme-linked immunosorbent
assay for the phomopsins is readily available to the food
industry. The most recent cultivars of L. angustifolius are resis-
tant to invasion by the responsible fungus (Diaporthe toxica), so
that the risk of phomopsin ingestion is now extremely low and
likely to be even less in the future.
Grain Processing
Dehulling
The simplest processing of lupin grain is repetitive abrasion of
the testa in a ripple flow mill, tangentially abrasive dehulling
device, or similar device. (The thickness and hardness of the
testa make it harder to dehull lupins than most grains.) Alter-
natively, the grain may be conditioned by the addition of
water, to swell the grain, followed by a mild heat treatment
to help break the bonding between the testa and cotyledons. It
is then possible to dehull using the same roller mills as used for
milling wheat and other cereals. The major components are
separated by air aspiration and grading across gravity tables.
The germ fraction usually separates with the hulls and is
mostly lost.
The main reason for dehulling is that the cotyledons are
much more nutritious than whole grain for nonruminant ani-
mals such as pigs, poultry, and fish. It is then possible to mill
and fiber-rich fractions for the feed, human food, and cosmetic
industries.
Milling
Hammer milling of whole grain to make a coarse meal is
common in the animal feed industry, simply to make the
constituents more available to ruminant animals by increasing
the surface area of material for digestion, and in some cases, it
may be uneconomic to dehull for nonruminant feeds. Ham-
70 mer cutting and roller milling of the splits are used to make a
2 range can be from 10 to 800 mm, which is far greater than for
cereal, pea, and soy flours.
<1
20 Fractionation
Several reports in the scientific literature and patents describe the
separation of protein, fiber, and oil fractions from cotyledons.
284 THE LEGUMES AND PSEUDOCEREALS | Lupin: Overview
Some processes cater for the recovery of alkaloids, for use as
biological agents against insect pests and some microorganisms,
and low-molecular-weight carbohydrates for use in the chemical
industry. These mostly involve wet grinding of the cotyledons
and separation of fractions according to solubility at different
pH or in different solvents. There may be an initial dry milling
and separation of particles by air classification.
Extrusion
Extrusion can improve the quality of lupin-based formulated
feeds. There is no expansion of the pellets as with starch-
enriched feeds, but there are improvements in feed digestibil-
ity, which makes the process cost-effective.
Food Uses
Traditional Uses
Lupins have been used as a food for as much as 6000 years in
the Andean highlands and over 3000 years around the Medi-
terranean. The pearl lupin of the Andean highlands of South
America, known locally as tarwi or chocho, was extensively
cultivated, and consumers would soak the seed in running
water to remove most of the bitter alkaloids before toasting
or boiling and drying to make kirku. Andean people have
long recognized the benefits of consuming lupins, associating
their use with religious festivals. In South America, the tradi-
tional use of tarwi as staple food continues in many Andean
communities. Around the Mediterranean, lupini beans, large-
seeded bitter grains of L. albus, have been extensively used as a
snack food and as a bean substitute in times of drought. They
are still commonly used as a snack bar. The beans (grains) are
substantially debittered before being pickled and bottled or
canned for use as a snack food. In the Middle East, the grain
of L. albus is used to make snack foods, most commonly at the
time of Ramadan (several thousand tonnes are imported from
Australia each year for this). They are usually boiled whole and
spiced, with the testa being removed before eating. Another use
is to make a paste similar to the traditional falafel from faba
beans.
Bean/Sprout
Immature seeds do not yet contain any alkaloids and have a
similar taste and nutritional value to immature (green) peas.
They can be used as an alternative to soybeans to make
edamame. Lupins can be germinated to make a sprout suit-
able for vegetable or salad use. Germination lowers not only
the content of alkaloids, phytate, and oligosaccharides on a dry
weight basis but also the overall content of the protein fraction.
The lupin sprout compares favorably with soy and mung
sprouts for taste and texture, although it has a slight beany
and bitter aftertaste. Unlike most species, there is no vitamin C
production on germination. There is, however, considerable
production of the isoflavones, genistein and daidzein.
Whole seeds may be deep-fried or made into spreads sim-
ilar in nature to hummus (traditionally made from chickpeas)
and falafel (traditionally made from faba beans) and used with
salads or breads.
Fermented Foods
Tempeh is a traditional Indonesian food, made by two suc-
cessive fermentations using soybeans as substrate. A bacterial
fermentation during the soaking of cooked dehulled soybeans
is followed by a solid-state fermentation of the bean mass by
the mold Rhizopus oligosporus. Tempeh has been successfully
made from L. albus, L. angustifolius, and blends of both with
soybeans. Indonesian consumers like the taste of lupin-based
tempeh but frequently comment that the texture is too firm.
The production of second-generation products, such as burgers
and patties, from lupin-based tempeh seems to overcome this
objection.
Miso is a fermented paste made from soybeans, usually
mixed with rice. A traditional process involves preparing a rice
koji by fermenting cooked rice with a culture of Aspergillus
oryzae (tane-koji) and then adding cooked soybeans and salt
to the koji. Lupins have been used experimentally to make
miso of comparable quality with soy miso. Natto, another
Japanese fermented food, can also be made from lupin grain.
Shoyu is the traditional soy sauce of Japan. Some Japanese
people now use lupin shoyu as an alternative. Lupins can be
used to make sauces similar in flavor and texture to the tradi-
tional soy sauces of China, Korea, and Indonesia.
Several thousand tonnes of ASLs have been used commer-
cially in Indonesia for tempeh production, and small commer-
cial batches of lupin-based miso have been sold in Japan.
Flour Additive
Lupin kernel flour can be mixed in with wheat or wholemeal
flours to make bread more nutritious, by giving a better bal-
ance of essential amino acids. Legume grains contain relatively
more lysine and less methionine than cereal grains; so, adding

10% lupin flour to wheat flours will give a product with
similar properties to the full wheat product but with an
improvement in amino acid score from under 40% to over
70% relative to egg albumin. Loaf volume is compromised
with increasing lupin content but fortifying with gluten or
using harder wheat in the blend can overcome this. In practice,
the limit of inclusion is
10%. There is an increase in water-
holding capacity. The texture, flavor, and golden color of the
lupinwheat flour are appealing to many consumers. In
Australia, some bread manufacturers use lupin hull flour to
provide bulk in high-fiber breads. Ultrafine kernel flour has
an attractive yellow color, good dispersion in aqueous systems,
and good emulsifying properties. Albus lupin flour can be
added into wheat flour to make the traditional Chilean breads
hallulla and marraqueta. Up to 10% micronized flour from
L. albus cotyledons could be included in breads, biscuits, and
cakes, enhancing protein quality, color, and taste. The added
lupin flour retards staling, probably by increased water reten-
tion and better emulsification properties. Studies have shown a
high acceptability of pasta enriched with L. angustifolius kernel
flour, and up to 15% substitution of L. albus flour for semolina
or durum flours enhanced the protein quality and appearance
of spaghetti noodles without affecting sensory qualities. Flour
from L. albus has been blended with noodle wheat flour to
make unsalted white noodles with improved protein and

dietary fiber contents and color and texture. Up to 50% lupin
flour can be incorporated into a range of cakes and biscuits.
There are several suppliers of lupin flour and other products
in Europe, but there are no available data on sales.
Lupin Kernel Fiber
The purified cell wall fraction from lupin kernels is virtually
colorless, odorless, and tasteless and can hold up to eight times
its own weight of water. The fiber has cholesterol-lowering
properties and acts as a fecal bulking agent, thus reducing
stool transit time and benefiting bowel health. It also enhances
satiety, probably due to its high water-holding capacity, and
has potential as an antiobesity agent. The soluble fiber fraction
can function as a fat replacer. The fiber can be used to supple-
ment breads, pastas, biscuits, mousses, and jellies.
Protein Fractions
Lupin kernel proteins have some valuable functional proper-
ties, combining good emulsifying, foaming, and water- and
oil-holding properties but with less thermal stability than soy
proteins. Nevertheless, lupin protein concentrates and isolates
are used as meat extenders and refined fractions used to
enhance the foaming properties of dairy desserts and other
foods.
Vegetable Milk Products
A lupin-based milk was used in the Chilean program for chil-
drens nutrition for several years and is still used by households
in some parts of the country.
In making tofu, it is possible to incorporate up to 30%
lupin (L. angustifolius) milk with soy milk before the coagula-
tion stage and produce an acceptable product, with the advan-
tage of a lower unit cost of production. Higher levels of
incorporation are not possible because the lupin proteins
lack the necessary tertiary structures to produce a good curd.
Sensory evaluations showed that tofu from a 30/70 lupin/soy
blend was equally acceptable as a tofu made from soy milk.
Fermentation of lupin milk produces a yogurt-like product
of stable texture and with no beany flavor. It is comparable
with the yogurts made from soy milk.
Malted lupins could be used as a health drink base. The
presence of the beneficial isoflavones genistein and daidzein
in the hypocotyls of developing lupin sprouts is well
reported. Lupin tea, made in the same way as pearl barley tea,
has been developed in Japan.
Allergenicity of Lupins
There are some reports of lupin allergenicity in the medical and
scientific literature. Nearly all of the affected subjects had a
history of reactions to other protein foods (e.g., peanuts, soy-
beans, and shellfish). The evidence to date suggests that lupin
proteins have a lower allergenicity than most other protein
foods.
THE LEGUMES AND PSEUDOCEREALS | Lupin: Overview 285
Feed Uses
Ruminants
Almost all of the testa can be fermented in the rumen, so there
is no need to dehull lupins for ruminant feeds. The whole grain
is highly fermentable with a minimal risk of acidosis. It is best
to crack the grain open as a minimum action, to increase access
for the rumen flora, but any grinding should not be too fine
because there can be a loss of material to the hindgut where it is
used less efficiently. A significant amount of protein degrada-
tion occurs in the rumen, but some protection from this is
possible by treating the grain with formaldehyde before feed-
ing. Other methods include flame roasting the grain, steaming
and rolling the grain into flakes, and coarsely grinding the
grain and then extruding the meal.
Lupins are a valuable component of diets of feedlot cattle
(up to 40% of the diet), and milking cows can readily consume
large amounts of (cracked) grain while in the milking sheds.
Rangeland farmers often supplement their grazing sheep with
50250 g grain per head per day in dry periods when pastures
are low. There is ample evidence for an enhancement of the
reproductive performance of both rams and ewes when their
diets are supplemented with lupin grain prior to mating. No
specific factor has been attributed to this.
Several hundred thousand tonnes of grain are retained on-
farm in Australia each year for grazing supplements for sheep.
The greatest consumption of traded lupin grain is by cattle,
either in feedlots or under intensive housing, when fed with
formulated diets containing lupin grain.
Nonruminant Animals
The testa is virtually worthless to all nonruminant species. It is
removed whenever economical to do so. Otherwise, it simply
supplies some bulk nonfermentable fiber. Processed lupin ker-
nel meal is frequently used in commercial feeds for pigs, poul-
try, and aquatic species. The actual inclusion levels will depend
on price and availability of the meal and alternative ingredients
and any constraints imposed by any individual ingredient. The
meal is low in lysine and methionine relative to most animal
needs, but this is not a significant problem. Firstly, this defi-
ciency is reflected in the price of the ingredient; secondly,
deficiencies of one ingredient in the formulation will be at
least partly offset by other ingredients; thirdly, crystalline
amino acids are readily available to the feeds industry.
Pigs
The digestible energy (DE) value of whole lupin grain for pigs
is
1415 MJ kg1; however, there is a lot of hindgut fermen-
tation and the net energy value is
10.5 MJ kg1. Nevertheless,
lupins are widely used in the pig industry. Recommended
inclusion levels of ASL vary from 100 to 350 g kg1 depending
on the age and physiological status of the animal. YLs have a
similar energy value and are also widely used. Albus lupins are
not used because they contain an inappetence factor for pigs.
This reduces feed intake, slows growth, and makes it uneco-
nomic to use such a diet.
The addition of industrial carbohydrase enzymes to lupin-
based pig diets can improve their net energy value.
286 THE LEGUMES AND PSEUDOCEREALS | Lupin: Overview
Pigs rank second to cattle in the overall consumption of
lupin grain, almost entirely as a component of formulated
feeds.
Poultry
The apparent metabolizable energy content of both ASL and YL
grains is
8.6 MJ kg1. It is not practical to use more than
100 g kg1 feed because of the high water-holding capacity of
the undigested carbohydrates, which can cause wet droppings.
For layer birds, it is possible to include up to 300 g kg1 in the
diets, as the wet droppings are less of a problem. Albus lupins
can be readily included in poultry diets. There is some benefit
from adding carbohydrase enzymes to poultry diets. The oli-
gosaccharides in lupin meals seem to have a beneficial effect on
the osmotic stability of the gut enhancing nutrient absorption.
Poultry ranks third as a consumer of lupins in formulated
feeds.
Finfish and crustaceans
Lupins can be used in both high nutrient density (HND; high
protein and oil content) and low nutrient density (low protein
and oil content) diets for finfish. When included in HND diets
fed to rainbow trout (Oncorhynchus mykiss), the DE value of
lupin kernel meals ranges from 12.8 MJ kg1 for L. angustifolius
to 14.8 MJ kg1 for L. albus. Protein digestibility of all lupin
varieties when fed to fish is high, typically greater than 95%.
The low essential amino acid content is less of a problem for
finfish fed with HND diets than for pigs and poultry because of
the high level of protein required in the diets restricting the
practical inclusion levels of the meal and also that so much of
the dietary protein is metabolized as an energy source. Good
acceptance and palatability of HND diets containing L. angu-
stifolius, L. albus, and L. luteus have been reported, with accept-
able inclusion levels up to 500 g kg1.
Lupin meals are used commercially in diets for freshwater,
estuarine, and marine finfish and crustaceans. This is a small
but emerging market, as the relative value of the splits seems
high for finfish.
Exercises for Revision
How can lupins be used in fermented foods?
What antinutrients can be present in lupins?
Characterize the major lupin proteins.
Exercises for Readers to Explore the Topic Further
Increase the applications of lupin flours in foods
Increase the industrial-scale production of traditional
lupin-containing foods
See also: Agronomy of Grain Growing: Lupin: Agronomy;
Breeding of Grains: Lupin: Breeding; Food Grains and the
Consumer: Grains and Health; Non-food Products from Grains:
Cereal Grains as Animal Feed; Processing of Grains: Extrusion
Technologies; Proteins: The Protein Chemistry of Dicotyledonous
Grains; The Basics: Grain: Morphology of Internal Structure; The
Legumes and Pseudocereals: Grain Legumes and Their Dietary
Impact: Overview.
Further Reading
Gladstones JS, Atkins C, and Hamblin J (eds.) (1998) Lupin as Crop Plants: Biology,
Production and Utilization. Oxon, UK: CAB International. This book is the definitive
work on all aspects of the Genus Lupinus, and was drawn on for this article.
Nahrung 45(6) (2002) A special edition covering the processing of lupins and other
grain legumes.
Petterson DS (2000) The use of lupins in feeding systems: Review. Asian-Australasian
Journal of Animal Sciences 13(6): 861882.
Van Barneveld RJ (1999) Understanding the nutritional chemistry of lupin (Lupinus sp.)
seed to improve livestock production efficiency. Nutrition Research Reviews
12: 130.
Wink M, Meissner C, and Witte L (1995) Patterns of quinolizidine alkaloids in 56
species of the genus Lupinus. Phytochemistry 38(1): 139153.
Relevant Websites
http://www.agric.wa.gov.au Department of Agriculture, Western Australia.
http://www.rala.is The International Lupin Association.
 Amaranth: Overview
H Corke and YZ Cai, Hubei University of Technology, Wuhan, China; University of Hong Kong, Hong Kong, China
HX Wu, Wuhan Chortle Biotechnology Company Limited, Wuhan, China
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The distribution and importance of different agricultural
crops can change fast: 80 years ago, soy bean was not
important in the United States; now, it is perhaps the
most valuable crop. Can grain amaranth rise from niche
status to major crop?
Grain Amaranthus, a small-seeded pseudocereal long culti-
vated in Central and South America, is fast-growing, adapt-
able, and generally pest- and disease-resistant.
Amaranthus produces a starchy seed with a nutritious non-
allergenic and gluten-free protein. The oil is high quality and
contains significant squalene. The plant can be used a source
of red pigments and as a leafy biomass for animal feed.
Research effort on Amaranthus lags far behind the countless
thousands of scientist-years devoted to major crops such as
maize, wheat, or rice. Hence, it is agronomically underde-
veloped and has modest grain yields. The market lacks a
critical mass of stable, uniform supply.
Modern biotechnology may aid the transfer of useful traits
from Amaranthus to other more established crop plants.
Learning Objectives
To understand the history and future potential of Amar-
anthus species as agricultural crops, for grain production,
for biomass, and for fractionation into value-added
components
To understand the properties and uses of Amaranthus
starch, protein, oil, pigments, and other fractions and how
these properties relate to new product development for
food and other applications
To understand the current state of Amaranthus production
and that in the major producer, China, its major use for
livestock feed
Amaranth Production and Development
Although amaranth has long been cultivated as a minor crop in
Central and South America and some areas of Asia and Africa,
its production seemed to rapidly expand from the early 1980s,
particularly in developing countries. However, growth seems
to have stagnated in recent years and no clear commercial
markets have developed. In many countries, especially in Asia
and Africa (e.g., China, India, Nepal, Ethiopia, and Kenya),
introduction, cultivation, and development of amaranth have
been undertaken. Since 1982, the Chinese Academy of Agricul-
tural Sciences has introduced hundreds of amaranth lines/
varieties from the United States and other countries to China.
Some selected superior varieties (e.g., R104 and K112) have
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00032-2
been successfully grown in over 20 provinces. Amaranth has
attained some popularity in China not only mainly for use in
feed but also for food and for other purposes. Although ama-
ranth is cultivated as a minor crop in China, it has been widely
used for animal feed because of its high biomass and good feed
value. At one time, amaranth production in China was claimed
to have reached 300 000 ha annually, ranking as the top ama-
ranth producer in the world. In the last 10 years, amaranth
production in China has not been changed greatly. It still ranks
on the top amaranth producer in the world. Some significant
production in Russia, India, and Mexico continues as well.
Despite the fact that in China and Russia there is larger-scale
cultivation of amaranth mainly for feed use, there is still appre-
ciable commercial cultivation of amaranth for human nutri-
tion in Mexico, South American countries, China, the United
States, Poland, Austria, etc. Current (2014) production in the
United States is estimated at 1200 ha, primarily to supply four
major corporate buyers (Arrowhead Mills, Health Valley, Ama-
ranth Resources, and Nu-World Amaranth).
There is considerable variation in amaranth yield per hectare
in different countries. In China, depending on environmental
conditions and cultivation systems, field production may yield
22005500 kg ha
1 of amaranth grain (Figure 1) and
90 000180 000 kg ha
1 of fresh plants (silage). During culti-
vation trials, 3000 kg ha
1 is achieved with good agronomic
practice and good seed sources, and even a maximum yield of
4000 kg ha
1 has been obtained in Montana, the United States,
and 6000 kg ha
1 in Peru. There was an average yield of
2250 kg ha
1 in the northwest hills of India and 2000 kg ha
1
in Austria. In the United States, extensive research work on
amaranth has been conducted, especially under an amaranth
research program at the Rodale Research Center, which started
in 1976 and lasted for 14 years. The richest genetic resources of
Amaranthus from many countries and regions in the world were
collected and introduced. Special breeding varieties and differ-
ent products have been developed, which are sometimes sold in
health food shops or supermarkets. Although amaranth culti-
vation in the United States ranges from the Great Plains to the
Rocky Mountains in Colorado, production is very limited, with
perhaps only a few hundred acres planted. Notably, through
collaboration of scientists from all over the world, initiated by
this center, amaranth development has been stimulated world-
wide. Research and development of grain amaranth in China;
amaranth biology, chemistry, and processing; and other
technologies have been summarized and reviewed. A series of
studies on Amaranthus were carried out at the University of
Hong Kong demonstrating the unusual properties of Amar-
anthus starch, protein, oil, and pigments and reveal their poten-
tial for use as food, being helpful to enhance future
development and utilization of amaranth. More recent notable
research is coming out of Argentina (e.g., papers by Anon et al.).
287
288 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview
Figure 1 Field production of grain amaranth in northern China,
illustrating the generally well-developed agronomic level leading to grain
production in the range of 4 tonne ha
1.
Biological and Agronomic Characteristics of Amaranth
Amaranth, a pseudocereal crop, is a dicotyledonous C4 plant
belonging to the genus Amaranthus (which consists of some 75
species) of the family Amaranthaceae. There are two major
types of amaranth: grain amaranth (e.g., A. cruentus L.,
A. caudatus L., A. hybridus L., and A. hypochondriacus L.) and
vegetable amaranth (e.g., A. tricolor L., A. dubius L., and
A. lividus L.). Grain amaranth is believed to originate from
Central and South America, whereas the main vegetable types
are believed to originate from Southeast Asia.
Amaranthus plants usually prefer hot, bright sunlight and
are distributed from the tropics to the semiarid regions. Culti-
vated species can grow in the tropics, subtropics, and temperate
zones and grow well even in some cold areas such as
Nepal and Chinese Tibet. Amaranth possesses high yield poten-
tial and high stress tolerance to drought, salinity, alkalinity,
or acidic soil conditions. It typically gives a grain yield of
22504500 kg ha
1 and a fresh leaf and stem weight of
30 00060 000 kg ha
1. The amaranth plant varies from
branched to unbranched, prostrate to erect, and dwarf to over
4 m in height. The leaves are normally elliptical, with an acute tip
and a cuneate base; leaf size varies significantly between and
within species. The flowers are indefinite inflorescences.
Amaranth seed morphology is unlike common true cereal
grains. Seed coat color ranges from black, brown, yellow, to
white. Seed embryo is campylotropous, that is, circular, with
the ends nearly touching and enclosing the perisperm
(Figure 2). The seeds are extremely small, with 0.51.2 g per
1000 seeds or 8501700 seeds per gram (3070 times smaller
than a typical wheat grain). A single plant can produce more
than 500 g of grain. However, in actual field production, seed
yield of amaranth varies greatly (1550 g per plant), depending
on the varieties used and soil fertility conditions.
Composition of Amaranth Grains and Vegetative
Tissues
Grains
Amaranth is one of the few multipurpose plants that can supply
grains and leafy vegetables of high nutritional quality for use as
food and feed. Amaranth grain is characterized by a relatively
higher content of protein, higher lipid content, and lower starch
content than those of the major cereals. A comparison of the
average chemical compositions between amaranth (A. hypochon-
driacus) and main cereals (maize, rice, and wheat) is shown in
Table 1. Table 2 shows the proximate chemical compositions of
the grains of several Amaranthus species, indicating that there is
some variation within an overall pattern among and within
species. Total dietary fiber content (soluble and insoluble) in
amaranth grains from A. caudatus, A. cruentus, and A. hypochon-
driacus ranges between 7.1% and 16.4%. The nonstarch
polysaccharides make up 67% in the light-colored or dark
grains. The content of various mineral elements and vitamins in
amaranth grains from major species was analyzed. The levels of P,
Ca, K, and Mg are usually higher than those found in common
cereal grains. The P/Ca ratio varies from 1.9 to 2.6. It is of
nutritional interest to find that amaranth grains contain relatively
higher level of iron than that in cereal grains. Amaranth grains are
relatively high in tocopherol (546 ppm), riboflavin, etc.
Leaves
Amaranth leaves contain 27.848.6% dry matter as crude pro-
tein and, on average, have somewhat more protein than spin-
ach, as shown in Table 3. The fat content ranges from 2.9% to
7.1% and contains linoleic acid as the major concentration of
unsaturated fatty acid (4055%) and palmitic acid as saturated
fatty acid (1825%). Amaranth leaves and stems are a very
good source of fiber (11.122.9%). Minerals such as P, Fe,
Mg, and Ca also exist in high concentrations, giving a total of
33.140.7% of ash content. Vitamins C and A are in concen-
trations of nutritionally significant levels, averaging 420 and
250 ppm, respectively. Seedlings, leaves, and stems of ama-
ranth extensively save as tasty vegetables and animal feeds in
the numerous places of the world, especially in China, India,
and Southeast Asia. The protein isolated from leaves of
Amaranthus species is rich in lysine (6.26.8%), higher than
that of alfalfa, cabbage, and maize. It also has relatively good
amounts of other amino acids. The edible and nutritional
quality of vegetable amaranth is almost equal to spinach.
In addition, it has been found that red-colored vegetative
tissues (seedlings, leaves, inflorescences, etc.) of Amaranthus spe-
cies contain high level of betalain pigments as natural colorants.
 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview 289

Cotyledons
Procambium
Starchy
Cotyledons perisperm

Seed Vascular
coat bundles

Endosperm
Shoot
apex
Perisperm
Procambium
Root

Procambium Radicle
(Root)
(a)
Cross-section Longitudinal section

(b)

Figure 2 (a) Illustration of A. cruentus seed in cross and longitudinal sections; (b) scanning electron micrograph of A. cruentus seed. Adapted
from Irving DW, Betschart AA, and Saunders RM (1981) Morphological studies on Amaranthus cruentus. Journal of Food Science 46(4): 1170; Becker R
(1994) Amaranth oil: Composition, processing, and nutritional qualities. In: Paredes-Lopez O (ed.) Amaranth: Biology, Chemistry, and Technology,
p. 135. Boca Raton, FL: CRC Press.

There has also been much information on other compounds in
amaranth (tannin, phytic acid, oxalate, saponins, nitrates, tryp-
sin inhibitor, etc.) with antiphysiological activity in animal
performance. Tannins are of importance from the nutritional
point of view, since they may influence protein digestibility and
reduce mineral bioavailability. Tannins in different amaranth
species range between 0.1% and 0.25% and seem to be low
enough not to have nutritional impact. The phytate content
(0.340.61%) in amaranth is higher than that found in rice
(0.10.14%), but lower than that detected in maize and wheat.
Amaranth Starch
Starch is the main component of amaranth grain, composing
4869% of its total dry weight. Starch generally consists of
straight-chain and branched-chain molecules, that is, amylose
and amylopectin. Table 4 shows that amylose makes up from
0% (waxy) to 34.3% of total starch in amaranth grain with the
balance being the amylopectin, showing that there is consid-
erable variation in amylose/amylopectin ratio among and
within the Amaranthus species. Light-colored grains normally
contain more starch (mean 69%) than dark-colored grains.
Starches isolated from A. hypochondriacus genotypes contain
the lowest amount of amylose (7.8%), whereas A. retroflexus
genotypes have the highest amylose (34.3%). The average
amylose content of 124 genotypes is 19.2%.
Amaranth starch granules come in polygonal, lenticular,
circular, and elliptical shapes. Amaranth starch granules are
extremely small, ranging from 0.8 to 2.5 mm in diameter. Com-
paratively, diameters of the granules in commercial starch range
from 3 to 8 mm for rice starch to 15100 mm for potato starch.
Amaranth starch granule size and its size distribution are char-
acteristics that influence the functional properties of starch.
290 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview

Table 1 Comparison of proximate composition between amaranth Since the 1970s, some interesting findings on physico-
grains and some cerealsa chemical and functional properties of amaranth starch have
been reported, for example, stable paste properties and a wide
Composition Amaranthb Wheat Corn Rice
range of viscosity, resistance to shear thinning, and gelatiniza-
Carbohydrate 59.2 66.9 67.7 75.4 tion temperatures. The focus of such research was restricted to
Crude protein 16.6c 14.0d 10.3e 8.5e very few Amaranthus species or genotypes and sometimes gave
Fat 7.2 2.1 4.5 2.1 contradictory results due to the genetic variation of the prop-
Crude fiber 4.1 2.6 2.3 0.9 erties of amaranth starch. A. hypochondriacus starch had a
Ash 3.3 1.9 1.4 1.4 higher amylograph viscosity, a higher gelatinization tempera-
Moisture 9.6 12.5 13.8 11.7 ture, a lower swelling power, and a higher solubility than
a wheat starch. A. paniculatus (synonym: A. cruentus) starch had
Percentage on dry weight basis.
b
Mean values of four Amaranthus species (A. cruentus, A. caudatus, A.
moderate swelling power, higher solubility, and lower pasting
hypochondriacus, and A. hybridus). viscosity than foxtail millet starch. Compared to maize starch,
c
N  5.85. A. paniculatus starch had a higher paste viscosity, lower paste
d
N  5.7. clarity, and higher freeze/thaw stability.
e
N  6.25. A large number of Amaranthus germplasm (26 species)
Data from Saunders RM and Becker R (1984) Amaranthus: A potential food and feed accessions were introduced into China for the evaluation of
resource. In: Pomeranz Y (ed.) Advances in Cereal Science and Technology, vol. 6, pp. amaranth starch properties. Compared to the reference maize,
357396. St. Paul, MN: American Association of Cereal Chemists; Singhal RS and rice, potato, and wheat starches, Amaranthus starch tended to
Kulkarni PR (1988) Composition of the seeds of some Amaranthus species. Journal of
have more stable paste, that is, lower pasting viscosities (lower
the Science of Food and Agriculture 42(4): 325331; Yue SX, Sun HL, and Tang DF
shear thinning and lower retrogradation), higher gelatiniza-
(1993) The Research and Development of Grain and Amaranth in China (in Chinese).
Beijing: Chinese Agricultural Science and Technology Press; Segura-Nieto M, Barba de
tion temperature, and higher energy of enthalpy for gelatiniza-
la Rosa AP, and Paredes-Lopez O (1994) Biochemistry of amaranth proteins. In: tion. Also, Amaranthus starch paste was more resistant to cold
Paredes-Lopez O (ed.) Amaranth: Biology, Chemistry, and Technology, pp. 75106. storage and had softer gel texture and lower changes of hard-
Boca Raton, FL: CRC Press. ness, cohesiveness, modulus, and adhesiveness. Figure 3

Table 2 Chemical composition of amaranth grains from various speciesa

Component A. caudatus A. cruentus A. hybridus A. hypochondriacus

Carbohydrate 59.662.8 60.762.6 58.6 57.0

Crude proteinb 17.618.4 13.218.2 14.0 17.9
Fat 6.98.1 6.38.1 6.7 7.7
Crude fiber 3.25.8 3.64.4 6.6 2.2
Ash 3.14.4 2.83.9 3.6 4.1
Moisture 9.511.6 6.28.8 10.5 11.1
a
Percentage on dry weight basis.
b
N  5.85.
Data from Singhal RS and Kulkarni PR (1988) Composition of the seeds of some Amaranthus species. Journal of the Science of Food and Agriculture 42(4): 325331; Yue SX, Sun
HL, and Tang DF (1993) The Research and Development of Grain Amaranth in China (in Chinese). Beijing: Chinese Agricultural Science and Technology Press; Segura-Nieto
M, Barba de la Rosa AP, and Paredes-Lopez O (1994) Biochemistry of amaranth proteins. In: Paredes-Lopez O (ed.) Amaranth: Biology, Chemistry, and Technology, pp. 75106. Boca
Raton, FL: CRC Press.

Table 3 Proximate composition of leaves from several Amaranthus speciesa

Amaranthus speciesb

Component 1 2 3 4 5 6 Spinach

Crude proteinc 48.6 46.5 33.3 27.8 44.5 32.7 34.4

Fat 3.3 6.8 2.9 3.9 4.6 7.1 3.2
Crude fiber 12.8 11.1 22.9 17.5 14.1 46.2
Ash 35.2 35.4 40.7 33.1 16.1
a
Percentage on dry weight basis.
b
(1) A. caudatus, (2) A. cruentus, (3) A. edulis, (4) A. hybridus, (5) A. spinosus, (6) A. tricolor.
c
N  6.25.
Adapted from Saunders RM and Becker R (1984) Amaranthus: A potential food and feed resource. In: Pomeranz Y (ed.) Advances in Cereal Science and Technology, vol. 6,
pp. 357396. St. Paul, MN: American Association of Cereal Chemists; Teutonico RA and Knorr D (1985) Amaranth: Composition, properties, and applications of a rediscovered
food crop. Food Technology 39(4):4954; Segura-Nieto M, Barba de la Rosa AP, and Paredes-Lopez O (1994) Biochemistry of amaranth proteins. In: Paredes-Lopez O (ed.)
Amaranth: Biology, Chemistry, and Technology, pp. 75106. Boca Raton, FL: CRC Press.
 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview 291

displays the Rapid Visco Analyzer (RVA) pasting profiles of
Amaranthus starch compared with maize starch. A wide range
of variation was found in the several tested properties of starch
among the Amaranthus species and among genotypes within
the same species, suggesting that the wide genetic diversity
necessitates specific choices for specific uses. Interestingly, it
was observed that there were significant effects of environment
on the tested properties of Amaranthus starch, indicating that
starch of the same genotype might have different starch prop-
erties if grown in different locations. Additionally, obvious
effects on pasting and rheological properties from genotypic,
pH, and salt factors were found.
Recently, amaranth starch has attracted increasing attention
because of its unusual properties. Possible product uses of
amaranth starch, capitalizing on its small granule size, could
include food thickeners, dusting powders for cosmetic as well
as food uses, laundry starch, and even possible biodegradable
Table 4 Starch and amylose content of amaranth grains (%)a
Species Starch
plastics. The unique properties of A. caudatus starch, namely, its
pasting properties and high susceptibility to amylase, might
represent potential uses as food, feed, and fuel. A. hypochon-
driacus starch had relatively low gelatinization temperature and
food freeze/thaw stability, indicating its use in gravies, sauces,
and soups. A. paniculatus starch could be used in salad dress-
ings and custard preparations. There were also reports on
chemical modification (e.g., succinylation and hydroxypropy-
lation) of amaranth starches and their properties.
Many studies demonstrated that starches isolated from
many Amaranthus species could be used as good thickeners
and stabilizers in salad dressing, canned food, sauce, frozen
food, etc., because of their stable paste, low retrogradation,
freeze/thaw resistance, and heating and cooling resistance.
Also, there has been a report that describes amaranth starch
as behaving very poorly in breads and cakes from the stand-
point of color and appearance, texture, and volume, probably
because of its low amylose content. During starch application,
it is worth paying attention to a fairly wide variation of ama-
ranth starch properties among different species and genotypes.
Amylose

A. cruentus 4863 22.1 (n 48)

A. hybridus 55 21.1 (n 14) Amaranth Protein
A. hypochondriacus 5262 7.8 (n 6)
A. retroflexus 34.3 (n 3) The second most abundant component of amaranth grains is
A. spinosus 18.1 (n 4) protein. Based on the amino acid composition, amaranth seed
A. tricolor 51 29.0 (n 8) protein is known to be of higher quality than most of the major
A. viridis 12.9 (n 3) cereal grains. The essential amino acid composition of ama-
124 genotypes 19.2  13.9
ranth grains from different Amaranthus species is summarized
Light-colored seeds 69  3 (n 9)
in Table 5. The lysine content is two to three times higher than
a
n, numbers of genotypes. that of the most common cereals, and sulfur amino acid con-
Data from Saunders RM and Becker R (1984) Amaranthus: A potential food and feed tent is also relatively high as compared with that of the most
resource. In: Pomeranz Y (ed.) Advances in Cereal Science and Technology, vol. 6, pp. important legumes (pea, bean, soybeans, etc.). Table 5 also
357396. St. Paul, MN: American Association of Cereal Chemists; Yue SX, Sun HL, and shows clear variation in amino acid composition among dif-
Tang DF (1993) The Research and Development of Grain Amaranth in China (in ferent species. Because amaranth protein has a better balance
Chinese). Beijing: Chinese Agricultural Science and Technology Press; Wu HX and of the essential amino acids than in cereals and legumes, it
Corke H (1999) Genetic diversity in physical properties of starch from a world collection
of Amaranthus. Cereal Chemistry 76: 877883.
Table 5 Essential amino acid composition of grains from three major
Amaranthus species (g per 100 g protein)
200 100
Profile Amino acid A. caudatus A. cruentus A. hypochondriacus
160 80 Cystine 2.3 2.03.8 2.03.9
Temperature (C)

K112
Viscosity (RVU)

Isoleucine 3.64.1 3.43.7 2.83.8

120 60 Leucine 5.96.3 4.85.9 5.05.8
Corn
Lysine 5.76.4 4.85.8 3.26.0
80 R104 40 Methionine 2.43.3 1.82.6 0.61.6
Phenylalanine 3.44.0 3.24.5 3.84.5
40 20 Threonine 3.8 3.24.2 2.64.3
Tryptophan 1.1 n.d. 1.14.0
Tyrosine 2.8 2.44.0 3.14.0
0 0
0 2 4 6 8 10 12 14 Valine 4.14.7 3.94.3 3.24.2
Time (min) MetCys 4.7 3.85.4 2.65.5
PhetTyr 6.2 5.68.5 6.98.5
Figure 3 RVA pasting profiles of two Amaranthus genotypes (K112 and
R104) compared with corn. Adapted from Wu HX, Yue SX, Sun HL, n.d., not determined.
and Corke H (1995) Physical properties of starch from two genotypes Data from Saunders RM and Becker R (1984) Amaranthus: a potential food and feed
of Amaranthus cruentus of agricultural significance in China. Starch/ resource. In: Pomeranz Y (ed.) Advances in Cereal Science and Technology, vol. 6, pp.
Starke 47: 295297; Corke H, Wu HX, Yue SX, and Sun HL (1997) 357396. St. Paul, MN: American Association of Cereal Chemists; Segura-Nieto M,
Developing specialty starches from new crops: A case study using grain Barba de la Rosa AP, and Paredes-Lopez O (1994) Biochemistry of amaranth proteins.
amaranth. In: Campbell GM, Webb C, and McKee SL (eds.) Cereals: In: Paredes-Lopez O (ed.) Amaranth: Biology, Chemistry, and Technology, pp. 75106.
Novel Uses and Processes, pp. 91102. New York: Plenum Press. Boca Raton, FL: CRC Press.
292 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview

scores higher than other seeds (e.g., wheat, barley, soybean, proteins showed similar effect, but to a much lesser extent.
and maize) in the FAO/WHO Nutritionists Protein Value Furthermore, amaranth protein concentrates were examined
Chart: a score of 100 is the ideal, whereas amaranth protein for direct application in certain food systems (wheat dough
received the highest score of 75, compared to cow milk (score and noodle). It was found that the mixing properties of wheat
72), soybeans (68), and peanuts (52), indicating that its bal- flour were improved by the addition of the protein concen-
ance was closer to the optimum required in the human diet trates. Addition of protein concentrates to a noodle formula-
(FAO/WHO). tion also showed some positive influence on final product
Seed proteins are generally classified into four types based quality. A few amaranth protein concentrates were also evalu-
on solubility: albumin, globulin, prolamin, and glutelin. Most ated as fillers for an emulsion-type meat product. Only one of
studies on amaranth proteins suggested that albumin was the the amaranth protein concentrates (genotype K112) gave some
major fraction (4965%), followed by glutelin (2242%), favorable results.
globulin (1418%), and prolamin (1.03.2%). However,
there was variation in amaranth protein fractions because of
protein extraction and fractionation procedures and due to Amaranth Oil
genetic variation. For instance, it was reported that the frac-
tionated proteins from five different Amaranthus species aver- Oil is another chemical component of much interest in ama-
aged albumin (65%), globulin (17%), prolamin (11%), and ranth grain, because it contains relatively higher levels of oil
glutelin (7%). Also, it was reported that the isolated proteins (58%) than cereal grains; moreover, there is a considerable
from A. hypochondriacus consisted of 4649% albumin plus amount of squalene in amaranth oil (5%) compared with that
globulin, 3% prolamin, and 3033% glutelin, and in another in oils of cereals and other oil crops (0.010.4%) (Table 7).
study of same species, similar percentages of albumin, globu- Palmitic, oleic, and linoleic acids are the major fatty acids in
lin, prolamin, and glutelin were 51.0, 15.9, 2.0, and 31.1, amaranth oil, with the saturated/unsaturated fatty acid (S/U)
respectively. ratios ranging from 0.12 to 0.50. Published data showed that
In the 1980s and 1990s, many researchers isolated major the fatty acid profiles in amaranth oil from different species
fractions (albumin, globulin, and glutelin) of proteins from were as follows: palmitic (1225%), oleic (1935%), linoleic
main Amaranthus species and examined their biochemical and (2562%), stearic (28.6%), and linolenic (0.32.2%) acids.
physicochemical properties and molecular weights. Based on The squalene levels range from 5% to 8% in A. cruentus oil.
their measurements of its emulsion activity, foaming stability, Lower amounts of squalene (25%) are tested in some
and surface hydroscopicity, along with its relatively high heat
stability, it was concluded that amaranth proteins could be Table 7 Oil and squalene content of amaranth grains and other seeds
useful as effective emulsifying and foaming agents. On the
other hand, there were reports that A. cruentus and A. hypochon- Species Oil (%) Squalene (%)
driacus protein fractions had poor emulsifying and foaming
A. cruentus (1) 5.577.72 4.525.44
properties, limited water absorption, and good gel-foaming A. cruentus (2)a 5.08.0
properties. A. hybridus 6.40 5.23
It was found that protein concentrates made from ama- A. hypochondriacus 5.357.05 3.625.01
ranth grains of five genotypes exhibited better solubility, emul- A. tricolor 5.08 6.14
sification, and foaming properties than the commercially Maize 4.0 0.03
available soy proteins (Table 6). The effect of proteins on Cottonseed 7.0 0.01
starch gelatinization and retrogradation was investigated Rice 1.03.0 0.3
using the hydrolyzed and native protein concentrates of ama- Olive 36.0 0.4
Peanut 47.0 0.03
ranth. The results suggested that native proteins tend to be
attracted toward the starch granule and contribute to the regu- a
Data from Lyon and Becker (1987), other data in the table from Becker R (1994) and He
lation of swelling and amylase leaching during gelatinization, HP, Cai YZ, Sun M, and Corke H (2002). Extraction and purification of squalene
therefore resulting in a weaker starch gel. However, hydrolyzed from Amaranthus grain. Journal of Agricultural and Food Chemistry 50: 368372.

Table 6 Functional properties of amaranth protein concentrations, compared to soy protein isolates

Species Genotypes Emulsifying activity (%) Solubility (%) Foam expansion (ml) Foam stability (ml)

A. cruentus K112 55.6 19.9 49 34

A. cruentus K350 73.3 56.5 91 56
A. cruentus K459 61.9 42.1 52 26
A. cruentus R104 56.0 47.2 35 18
A. hybridus No.3 62.2 60.5 68 26
Soybean Soy A 50.6 33.3 23 20
Soybean Soy B 45.7 21.3 28 22
LSD0.05a 3.1 4.5 20 9
a
Least significant difference (p < 0.05) for comparison of means in the same column.
Data from Bejosano FP and Corke H (1999) Protein quality evaluation of Amaranthus wholemeal flours and protein concentrates. Industrial Crops and Products 76: 100106.
 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview
Amaranthus species of West Africa. Other lipids in amaranth,
such as triglycerides, sterols, methyl sterols, terpenic and ali-
phatic alcohols, tocopherols, and hydrocarbons, have also
been identified. Recent results indicated that the major fatty
acids in amaranth oil from 11 Amaranthus genotypes included
palmitic acid (19.123.4%), oleic acid (18.738.9%), and
linoleic acid (36.755.9%), with S/U ratios of 0.260.32. The
squalene content in amaranth oils from 11 genotypes of four
species ranges from 3.6% to 6.1% (Table 7). These reports
reveal the significant variation in oil composition content
among various species and genotypes.
Squalene is a biosynthetic precursor to all steroids. It is an
important functional ingredient in skin cosmetics and also as an
excellent computer-disk lubricant. It has been reported that the
decreased risk for some cancers might be associated with squalene
consumption, and the use of squalene alone is effective in decreas-
ing serum cholesterol levels. The traditional source of squalene is
from shark and whale liver oil. Estimates in 2015 place the world
market at 2600 tonne year
1, with a growth in demand of per-
haps 10% per year. It is very expensive ($43 000 per tonne).
Amaranth oil will attract more attention as an alternative plant
source of squalene because of its abundant squalene content,
although efforts to further increase the level are necessary.
Squalene can be easily extracted from amaranth grain by
simple vacuum distillation. Recently, work to screen genotypes
containing a higher content of squalene in a wide range
of Amaranthus species and to develop efficient large-scale
separation technique has been conducted; a method was estab-
lished to separate and purify squalene from amaranth oils.
After saponification, squalene content increased from 4.2%
to 43.3% in the unsaponifiables by the removal of the saponifi-
ables. The unsaponifiables were fractionated by a chromato-
graphic method to obtain highly purified squalene. The final
purity of squalene reached 9498%.
However, amaranth oil occurs in very low levels and at
higher cost than oils of peanut, olive, rapeseed, etc. Unless a
specialty starch or protein could also be produced from the
defatted meal, a substantial decrease in raw product cost will
probably be required before amaranth oil goes commercial.
Amaranth Pigments
There has been an increasing trend toward replacement of
synthetic colorants with natural pigments since the 1980s
Table 8 Pigment content and fresh weight of 12 genotypes from seven Amaranthus species
Species Genotypes Fresh weight of colored parts (kg ha
1)a
A. cruentus 12 56 390  7225 (Lv, In)
A. caudatus 2 42 520 (In)
A. tricolor 3 20 520 (Lv)
A. hypochondriacus 1 29 160 (In)
A. hybridus 1 39 285 (In)
A. lividus 1 17 370 (Sl)
A. paniculatus 1 44 550 (In)
FW, fresh weight.
a
Sl, seedlings; Lv, leaves; In, inflorescences. Estimated as Sl, Lv, or In weight per plant  planting density.
Data from Cai YZ, Sun M, Wu HX, Huang RH, and Corke H (1998) Characterization and quantification of betacyanin pigments from diverse Amaranthus species. Journal of Agricultural
and Food Chemistry 46(6): 20632070.
293
because of safety and health concerns. Red-colored vegetative
tissues of Amaranthus plants contain a high concentration of
betalain pigments and are often produced in high biomass
(Table 8). They have attracted considerable interest as potential
substitutes for the well-known betalains from beetroot (Beta
vulgaris), which are extensively used in the food industry world-
wide. A comprehensive systematic study on amaranth pigments
has covered field evaluation and selection of betalain-
producing genotypes/species; extraction, identification, and
quantification of betalain components; determination and
evaluation of properties and antioxidant activity; and pigment
production and utilization in certain food systems.
Betalains are divided into two major structural groups, that
is, red-violet betacyanins and yellow betaxanthins. Amaranth
pigments belong to betacyanins. They could be easily extracted
with water or methanol. Sixteen kinds of betacyanins were
isolated and characterized from plants in the Amaranthaceae.
Figure 4 shows HPLC profile of amaranthine-type betacyanins
and their molecular structures, which are predominant pig-
ments in all Amaranthus species. Quantitative analysis data
showed that the screened amaranth species/varieties yield
higher betacyanin pigments than feasible in beetroot produc-
tion. Colorant properties and stability of the betacyanins are
the key factors of pigment quality that directly influence their
application in food systems. Assessment of color characteristics
and stability of betacyanins from Amaranthus species showed
that Amaranthus betacyanins exhibited bright red-violet color
and favorable stability under selected conditions (pH 57 and
temperature <14  C), as compared to red radish anthocyanins.
Dried Amaranthus pigments had good storage stability (t1/
2 23.3 months) at room temperature. The presented data
support the potential development and utilization of Amar-
anthus betacyanins as new natural colorants for use in the food
industry, particularly for low-temperature uses.
Aqueous extracts of natural pigments are commonly
unstable and sensitive to storage environmental factors. Spray
drying can significantly improve storage stability. Amaranthus
betacyanin extracts were spray-dried using a range of 1025 DE
maltodextrins as carrier and coating agents at different
inlet/outlet air temperatures and various feed solid contents.
The best dried pigment powder made was superior to commer-
cial red beet powder in terms of physical properties (Table 9).
The results demonstrated the feasibility of the production
of spray-dried Amaranthus betacyanin extracts as a food-grade
colorant.
Total betalain content (mg per 100 g FW)
42199
82144
51143
47
82
46
127
294 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview

mAU
1
1: amaranthine HOCH2
2500 HO O
1: isoamaranthine (C15 6 1 O H
epimer) HO 5 2
HOOC 7 + COO
2000 O O HO N
HO
2
HO 1
1500 OH
1 H 15
1000 N COOH
HOOC
H
500

0 2.5 5 7.5 10 12.5 15 17.5 20 min

Figure 4 Structure and HPLC elution profile of amaranthine-type betacyanins from A. tricolor. The isomeric form (10) is the C-15 epimer of
corresponding betacyanin (not drawn).

Table 9 Comparison of amaranth pigment powder and commercial beet powder (No. 3600 or E162)a

Color parameters
Spray-dried Betacyanin content Hygroscopic moisture Pigment
powder (%)b (g/100 g) L* a* b* H0 retention (%)c

Amaranth powder 0.74 48.5 49.35 32.60
2.47 355.7 87.9  1.8
(
4.3)
Beet powder 0.31 58.8 51.54 19.64 0.71 2.0 84.3  1.4
a
Amaranth pigment powder made by spray drying at 180C inlet temperature using 19.8% solids in feed mixture (15 DE maltodextrin).
b
Calculated as % amaranthine and % betanin.
c
Determined at 25  C and 32% RH after 16 week storage.
Data from Cai YZ and Corke H (2000) Production and properties of spray-dried Amaranthus betacyanin pigments. Journal of Food Science 65: 12481252.

Application of Amaranthus betacyanins in jelly, ice cream,
and a model beverage was evaluated by comparison to com-
mercial colorants (red radish anthocyanins and synthetic
colorant FD&C Red#3). The results revealed the feasibility
and potential of Amaranthus betacyanins for use in the model
food systems at low temperature. Amaranthus betacyanins in
the foods examined exhibited better color characteristics than
red radish anthocyanins at the same levels, but were not as
bright as FD&C Red#3. The betacyanins had color stability
comparable to red radish and FD&C Red#3 at lower tempera-
ture (<14  C) and also similar color stability at 25  C, but not
as stable as the two commercial colorants at higher tempera-
ture (37  C). Additionally, all betalains isolated from plants in
the Amaranthaceae exhibited strong antioxidant activity. To
date, there has been evidence suggesting that certain natural
colorants as nutritional antioxidants in the diet may reduce the
risk of some diseases. This gives the possibility of these beta-
lains being used both as natural colorants and as potential
natural antioxidants.
Food and Feed Utilization of Amaranth
Traditional foods are made from amaranth grains in many
countries, for example, alegria and atole in Mexico,
alboroto in Guatemala, bollos in Peru, Chapati in
Himalayas, laddoos in India, and sattoo in Nepal. These
amaranth-based foods are still consumed today in various
areas of the world. Modern food applications of amaranth
grains include breakfast foods, infant/weaning food formula-
tions, breads, pastas, flakes, drinks, and beverages. Currently
available consumer food products include popped foods, bak-
ing mixes, malted flours, breakfast cereals, snacks, baked
goods, and whole amaranth products. In China, a wide range
of amaranth food products has been developed for retail sale.
These include instant flour for beverages or porridge, packaged
dried noodles, cakes, biscuits, and soy sauce made from
brown/black amaranth and other grains (wheat and soybean).
China has a wide diversity of traditional food products made
from specialty grains, such as buckwheat, millets, sorghum,
and legumes. By adapting products made from these, addi-
tional amaranth-based foods have been developed, such as
composite flour instant noodles, distilled spirits, and vinegar.
Based on the properties of diverse fractions of amaranth grain,
and depending on the physical, nutritional, and sensory
requirements of specific products, there is continuing potential
for new product development based on amaranth. Examples of
recent studies include that of Martinez et al. who investigated
the effect of A. mantegazzianus flour on the technological and
sensory quality of bread wheat pasta. Also, Diaz et al. drew
attention to the significance of amaranth as source of gluten-
free ingredients. They added 20% amaranth to a maize base
 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview
used in twin screw extrusion. Addition of amaranth conferred
increased sectional expansion index. Also, in recent years, ama-
ranth grains have been developed and processed for many
innovative new food products in China, such as amaranth-
flavored cake, amaranth yogurt, and amaranth Taosu (a type
of traditional Chinese sweet cake).
Furthermore, vegetable amaranth has been used as a tradi-
tional vegetable for several centuries in China. In sub-Saharan
Africa and Central and South America, vegetable amaranth is
also an important leafy vegetable, consumed as a cooked green
vegetable or used in soup.
Amaranth is a fast-growing, high-nitrogen, and high-
biomass plant that can be used fresh or as silage for animal
feed. Amaranth grain flour or powdered leaf meal can be used
to make commercial pelleted or other compound feeds. Cur-
rently, feed application of amaranth has been successful and
popular in China and has played a role in developing Chinese
animal husbandry. Amaranth feeds made from leaves, stems,
panicles, and grains include green fodder, silage, leaf and seed-
meal feed, pellet feed, etc. The evaluation of feed use has
shown that amaranth feeds efficiently raise pigs, chickens,
cattle, sheep, and fish to improve the quality and productivity
of animal production. Chinese peasant households are accus-
tomed to raise livestock and poultry, and nowadays, in many
areas, amaranth is the main source of silage for household pig
production. However, it is not widely used as feed in other
countries.
Other Uses
Considerable further research has been conducted over the
past 10 years on potentially healthful or functional compo-
nents of amaranth grain. Amaranth grain was 11% dietary
fiber, of which 78% was insoluble and 22% was soluble.
These levels were similar to quinoa, but the soluble fraction
was greater than in maize. Quiroga et al. tested lectins from the
amaranth protein hydrophobic fraction of A. mantegazzianus
and A. caudatus and found high activity as cell-proliferation
inhibitors, the effect being attributed to a cytotoxic effect,
followed by apoptosis, and inhibition of cell adhesion. Simi-
larly, Sabbione et al. produced fractions and hydrolysates of
amaranth seed protein fractions and showed that antithrom-
botic activity was particularly high in the glutelin fraction. In a
general review of amaranth nutritional components, Ven-
skutonis and Kraujalis summarized the potentially valuable
fractions of the plant, classifying into macrocomponents
(lipids, proteins, and carbohydrates including dietary fiber
and starch) and other important constituents, such as squa-
lene, tocopherols, phenolic compounds, phytates, and
vitamins.
Conclusions
In summary, amaranth possesses high yield potential and high
stress tolerance to drought, salinity, alkalinity, or acidic soil
conditions. Its grains have attractive chemical composition,
superior nutritive value, and some special functional proper-
ties. Its vegetative parts with high biomass can be used for
295
animal feed. Its red-colored vegetative tissue with high level
of betacyanins is a potential alternative source of well-known
beetroot pigments (betalains). Amaranth has been attracting
worldwide attention and will be a high-potential new crop
with multiple uses. Since the first edition of this article 12
years ago, more than 250 research papers in international
scientific journals have been published on food science aspects
of amaranth, most regarding novel applications of nutritional
fractions of the grain. This developing corps of technical
knowledge may help to increase demand to a level that will
justify investment in major production and processing sys-
tems. However, intense and continuous efforts are necessary
in several aspects, such as amaranth breeding and field culti-
vation (for yield, shattering, and lodging), relevant food/feed
processing and development, product industrialization, and
marketing.
Exercises for Revision
What makes Amaranthus most suited for food or for feed
production?
What are the general properties of Amaranthus starch and
protein?
Exercises for Readers to Explore the Topic Further
For your production conditions and regulatory environ-
ment, is Amaranthus betacyanin a viable competitor for
red beet pigments? Consider potential yield, production
cost, coproducts, and range of pigment colors that can be
produced.
Squalene is a high-value lipid, with critical applications in
the computer industry. Animal sources are limited and
unsustainable, and plant sources such as Amaranthus yield
low amounts relative to total biomass. What is the potential
for genetic engineering of microbes or other high-yield
crops with Amaranthus genes for enhanced squalene
production?.
Speculate or brainstorm on what it would take to turn
Amaranthus into a breakthrough crop leading to production
at the level of maize or soy bean within 50 years.
See also: The Cereal Grains: Coix: Overview; The Legumes and
Pseudocereals: Buckwheat: Overview; Pseudocereals: Overview;
Quinoa: Overview.
Further Reading
AgMRC (2015) http://www.agmrc.org/commodities__products/specialty_crops/
amaranth/ Agricultural Marketing Resource Center, Iowa State University.
Becker R (1994) Amaranth oil: Composition, processing, and nutritional qualities.
In: Paredes-Lopez O (ed.) Amaranth: Biology, Chemistry, and Technology,
pp. 133141. Boca Raton, FL: CRC Press.
Bejosano FP and Corke H (1999) Protein quality evaluation of Amaranthus wholemeal
flours and protein concentrates. Industrial Crops and Products 76: 100106.
Berghofer E and Schoenlechner R (2002) Grain amaranth. In: Belton PS and Taylor JRN
(eds.) Pseudocereals and Less Common Cereals Grain Properties and Utilization
Potential, pp. 219260. Berlin: Springer.
296 THE LEGUMES AND PSEUDOCEREALS | Amaranth: Overview
Cai YZ and Corke H (2000) Production and properties of spray-dried Amaranthus
betacyanin pigments. Journal of Food Science 65: 12481252.
Cai Y, Sun M, and Corke H (2005) Development and application of betalains from the
Amaranthaceae. Trends in Food Science and Technology 16: 370376.
Cai YZ, Sun M, Wu HX, Huang RH, and Corke H (1998) Characterization and
quantification of betacyanin pigments from diverse Amaranthus species. Journal of
Agricultural and Food Chemistry 46(6): 20632070.
Chen JZ (2011) Development and preparation of amaranth yogurt. Food Research and
Development (in Chinese) 12: 7273.
Corke H, Wu HX, Yue SX, and Sun HL (1997) Developing specialty starches from new
crops: A case study using grain amaranth. In: Campbell GM, Webb C, and
McKee SL (eds.) Cereals: Novel Uses and Processes, pp. 91102. New York:
Plenum.
Diaz JMR, Kirjoranta S, Tenitz S, et al. (2013) Use of amaranth, quinoa and kaniwa in
extruded corn-based snacks. Journal of Cereal Science 58: 5967.
He HP, Cai YZ, Sun M, and Corke H (2002) Extraction and purification of
squalene from Amaranthus grain. Journal of Agricultural and Food Chemistry
50: 368372.
Irving DW, Betschart AA, and Saunders RM (1981) Morphological studies on
Amaranthus cruentus. Journal of Food Science 46(4): 11701174.
Lamothe LM, Srichuwong S, Reuhs BL, and Hamaker BR (2015) Quinoa
(Chenopodium quinoa W.) and amaranth (Amaranthus caudatus L.) provide
dietary fibres high in pectic substances and xyloglucans. Food Chemistry
167: 490496.
Leon-Camacho M, Garcia-Gonzalez DL, and Aparicio R (2001) A detailed and
comprehensive study of amaranth (Amaranthus cruentus L.) oil fatty profile.
European Food Research and Technology 213: 349355.
Lyon CK and Becker R (1987) Extraction and refining of oil from amaranth seed. Journal
of the American Oil Chemists Society 64: 233236.
Martinez CS, Ribotta PD, Anon MC, and Leon AE (2014) Effect of amaranth flour
(Amaranthus mantegazzianus) on the technological and sensory quality of bread
wheat pasta. Food Science and Technology International 20: 127135.
Paredes-Lopez O (1994) Amaranth: Biology, Chemistry, and Technology. Boca Raton,
FL: CRC Press.
Quiroga AV, Barrio DA, and Anon MC (2015) Amaranth lectin
presents potential antitumor properties. LWT Food Science and Technology
60: 478485.
Sabbione AC, Scilingo A, and Anon MC (2015) Potential antithrombotic activity
detected in amaranth proteins and its hydrolysates. LWT Food Science and
Technology 60: 171177.
Saunders RM and Becker R (1984) Amaranthus: A potential food and feed resource.
In: Pomeranz Y (ed.) Advances in Cereal Science and Technology, vol. 6,
pp. 357396. St. Paul, MN: American Association of Cereal Chemists.
Segura-Nieto M, Barba de la Rosa AP, and Paredes-Lopez O (1994) Biochemistry of
amaranth proteins. In: Paredes-Lopez O (ed.) Amaranth: Biology, Chemistry, and
Technology, pp. 75106. Boca Raton, FL: CRC Press.
Singhal RS and Kulkarni PR (1988) Composition of the seeds of some Amaranthus
species. Journal of the Science of Food and Agriculture 42(4): 325331.
Sleugh BB, Moore KJ, Brummer EC, Knapp AD, Russell J, and Gibson L (2001) Forage
nutritive value of various amaranth species at different harvest dates. Crop Science
41: 466472.
Teutonico RA and Knorr D (1985) Amaranth: Composition, properties, and applications
of a rediscovered food crop. Food Technology 39(4): 4954.
Venskutonis PR and Kraujalis P (2013) Nutritional components of amaranth seeds and
vegetables: A review on composition, properties, and uses. Comprehensive Reviews
in Food Science and Food Safety 12: 381412.
Wang YY, Wang HS, Zhang HT, and Cui YH (2011) Research advances of amaranth as a
high-quality feed plant. Feed Research (in Chinese) 1: 1920.
Williams JT and Brenner D (1995) Grain amaranth (Amaranthus species).
In: Williams JT (ed.) Cereals and Pseudocereals, pp. 129186. London: Chapman
and Hall.
Wu HX and Corke H (1999) Genetic diversity in physical properties of starch from a
world collection of Amaranthus. Cereal Chemistry 76: 877883.
Wu HX, Yue SX, Sun HL, and Corke H (1995) Physical properties of starch from two
genotypes of Amaranthus cruentus of agricultural significance in China. Starch/
Starke 47: 295297.
Yue SX, Sun HL, and Tang DF (1993) The Research and Development of Grain
Amaranth in China (in Chinese). Beijing: Chinese Agricultural Science and
Technology Press.
Zhang FF, Jiang YP, Cao YJ, and Sun HN (2014) Feed value of amaranth and its
utilization in poultry. Poultry Science (in Chinese) 3: 68.
Relevant Websites
http://www.agmrc.org/commodities__products/specialty_crops/amaranth/ Website
of the Agricultural Marketing Resource Center, Iowa State University.
 Beans: Overview
SK Sathe, Florida State University, Tallahassee, FL, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
On a caloric basis, dry beans are more nutrient-dense than
cereals.
Among dry beans, the common bean Phaseolus vulgaris is
globally consumed in the largest quantity.
Dry beans contain two to three times more protein than
cereals.
Dry beans are low in lipids and contain no cholesterol.
Dry beans are a good source of complex carbohydrates
including fiber.
Dry beans contain several different types of proteins that
include storage proteins (e.g., 7S and 11S).
Dry beans contain antinutritional factors.
Dry beans are consumed in a variety of forms including
split cotyledons (dal), sprouts, bean flour, porridge,
fermented beans, gruel, fried beans, and others.
Dry beans are often used with cereals and/or animal foods
(e.g., meats).
Learning Objective
To achieve understanding of the global importance of
legume plants, particularly the dry beans, in human food
and animal feed supply
Introduction
Globally, of the 50 000 edible plant species, 15 crop plants
provide 90% of the worlds food energy intake. Of these 15
crops, three cereals rice, maize, and wheat make up two-
thirds of this 90% of the worlds food energy intake as they serve
as the staples for over 4000 million people. Comparatively,
contribution of legumes is less. However, because legumes
contain almost two to three times more protein than cereals,
their dietary importance as protein source is well appreciated.
Of more than 1300 species of legumes, only
20 are most
commonly consumed by humans. Among these, the common
dry bean, Phaseolus vulgaris, is consumed in the largest quantity
on a worldwide basis. Dry beans (excluding oilseeds) are low in
lipids, are low in sodium, and contain no cholesterol. They are a
rich source of proteins, complex carbohydrates including fiber,
vitamins, and certain minerals. On a caloric basis, dry beans are
more nutrient-dense than cereals. Dry beans are less expensive
than animal food products and, when stored properly, have
considerably longer shelf life than several animal, fruit, and
vegetable products. Since legumes have the ability to fix atmo-
spheric nitrogen and therefore add nitrogen to the cropsoil
ecosystem, they are important in soil conservation and mainte-
nance of soil quality.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00033-4
Global Distribution, Varieties, and Commercial
Importance
The word legume is derived from the Latin word legumen,
which means seeds harvested in pods. The term pulse (from
Latin word puls meaning pottage) is used for legume seeds
that contain small amounts of fat, while for those containing
large amounts of fat (such as soybeans and peanuts), the term
leguminous oilseed is used. According to the Food and Agri-
culture Organization (FAO), the word legume is used for all
leguminous plants. The most commonly used legumes as
human food are listed in Table 1.
Although legumes have been cultivated for several thou-
sand years, the chronology and origins of domestication of
food legumes are almost impossible to reconstruct. Some
legumes (e.g., lentils) have been dated back to 70006000
BC. Leguminosae (or Fabaceae) is the third largest family of
flowering plants (after Compositae and Orchidaceae) in size
and economic importance and is second only to the grasses
(Gramineae). Current estimates indicate that Leguminosae has

16 00019 000 species in
750 genera. The subclassification
is somewhat controversial. Almost all of the domesticated
legumes used as food are members of Papilionoideae. All of
the common beans belong to the tribe Phaseolae.
Global production of dry beans and pulses (Table 2)
remains important for human and animal nutrition. On a
worldwide basis, the common beans (Phaseolus spp.) are the
number one crop among dry beans (excluding oilseeds) in
both production and consumption and are therefore econom-
ically an important crop. In 2009, the global dry bean supply
was 2.4 kg per capita per year. Asia produces the largest quan-
tity (46.11% of the total world production) of dry beans fol-
lowed by the Americas (31.08%), Africa (20.48%), Europe
(2.11%), and Oceania (0.22%). In 2011, Brazil (12.19%),
India (15.69%), China (6.32%), Myanmar (16.08%), Mexico
(4.67%), and the United States (6.26%) accounted for 61.21%
of the global dry bean production. Europe (21.41%), the
Americas (16.08%), and Africa (10.28%) lead world imports
of dry beans in terms of dollar value (Table 2). India, Japan,
the United States, Brazil, the United Kingdom, Italy, Mexico,
and China, respectively, accounted for 17.58%, 6.70%, 6.13%,
4.60%, 4.35%, 4.22%, 3.18%, and 2.07% of the total dollar
value (42.76%) of worldwide imports ($3 227 563 000) of dry
beans. Asia (49.55%) and the Americas (34.99%) account for
84.54% of the global export of dry beans (Table 2). In 2011,
the leading exporting countries of dry beans were China,
Myanmar, Argentina, the United States, and Canada, respec-
tively, accounting for 29.17%, 15.88%, 10.67%, 9.74%, and
6.96% of the total dollar value (72.42%) of worldwide dry
bean exports.
297
298 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview

Table 1 Grain legume species commonly used for food purposes

Botanical name Common name

Arachis hypogaea Groundnut, peanut, monkey nut, goober pea, nguba

Cajanus cajan Pigeon pea, arhar, red gram, tur, toovar Angola pea, gandal, ambre vade, alverja
Ca. indicus Pigeon pea, Congo pea, yellow dal
Canavalia ensiformis Jack bean, horsebean, gotani bean, haba de burro, Chickasaw lima
C. gladiata Sword bean, maxima
Cicer arietinum Chickpea, Bengal gram, chana, desi chana, kabuli, chiche
Ci. minotinum Chana, garbanzo
Cyamopsis tetragonoloba Cluster bean, guar, aconite, cyamopse
Dolichos biflorus Horse gram
D. lablab Hyacinth bean, bonavist, field bean, caballeros, Indian butter bean, val bean, Egyptian kidney bean
Ervum vulgaris Lentils, masoor dal
Faba vulgaris Windsor bean
Glycine max Soybean, soja
G. hispida
G. soja
Lablab niger Lablab bean
La. purpureus Kidney bean, hyacinth bean, Indian bean, lubia bean
Lathyrus sativus Grass pea, kesari dal, vetch, chickling vetch, chicaro
Lens esculenta Lentils, masoor dal, red dal, lentil, split pea, lentija
Le. culinaris
Lupinus spp. Lupins, tarwi, tarin, pearl lupin, wolf bean, tremoco
Macrotyloma uniflorum Horse gram, Madras gram, kallu, kulthi bean
Mucuna pruriens Velvet bean, cowage, Mauritius bean, stizolobia
Phaseolus aconitifolius Moth bean
Ph. acutifolius Tepary bean, pavi, Yorimuni, dinawa
Ph. angularis Adzuki bean, feijao
Ph. aureus Mung bean, green gram, golden gram, chiroko, chicka sano pea
Ph. calcaratus Rice bean, frijol arroz
Ph. lunatus Lima bean, sieva bean, Madagascar bean, sugar bean, Burmabean, towe bean, pole bean, caraota, panguita
Ph. mungo Mung bean, mungo bean, urd dal, black gram, urad, woolly pyrol, kambulu
Ph. radiatus Mung bean, golden graham, green gram
Ph. vulgaris Dry bean, haricot, common bean, kidney bean, navy bean, pinto or snap bean, feijao, opoca, rajma, French bean,
chumbinho
Pisum sativum Dry pea, green pea, garden pea, field pea
Pi. angularis
Pi. arvense
Psophocarpus Winged bean (humid tropics), goa bean, asparagus bean, Colombo, four-angled bean, princess bean
tetragonolobus
Sphenostylis stenocarpa Yam bean
Stizolobium spp. Velvet bean
Tetragonolobus purpureus Winged bean (Europe)
Trigonella foenumgraecum Methi, fenugreek
Tylosema esculentum Marama bean
Vicia faba Broad bean, horsebean, faba bean, field bean, Windsor bean
Vic. sativa Vetch
Vigna aconitifolia Moth bean, matki, mouth bean, mat, math
Vig. aureus Mung bean
Vig. radiata
Vig. mungo Black gram, urd, urad, kambulu, pyrol
Vig. sinensis Dry cowpea
Vig. umbellata Rice bean, red bean, mambi bean
Vig. unguiculata Black-eyed cowpea, black-eye pea, cowpea, kaffir bean, Hindu pea, asparagus pea
Voandzeia subterranea Bambara groundnut, Madagascar groundnut, earthpea, Congo goober, kaffir pea, jugo bean, haricot pistache

Data from Deshpande SS and Srinivasan D (1990) Food legumes: Chemistry and technology. Advances in Cereal Sciences and Technology 10: 147241; Doughty J and Walker A
(1982) Legumes in Human Nutrition. Food and Nutrition Paper 20, 152 pp. Rome, Italy: Food and Agriculture Organization of the United Nations.

Morphology of the Pods/Seeds the seed coat, the cotyledons, and the embryo axis. In most dry
seeds, they account for 820%, 8090%, and 12% of the seed
Regardless of the fat content, most legume seeds have similar weight, respectively. The majority of the nutrients are present
structure. Mature legume seeds have three major components: in cotyledons. Typical seed structure and various anatomical
 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview 299

Table 2 Acreage, production, yield, and import/export data for pulses and dry beans

Area harvested (Ha) Yield (Hg/Ha) Production (tons) Import Export

Tons $ Tons $
P D P D P D D D D D

World 5 903 158 28 780 377 8554 8040 5 049 539 23 140 276 3 321 726 3 227 563 000 3 393 895 3 074 549 000
Africa 1 890 425 7 198 019 6357 6585 1 201 768 4 740 016 408 551 331 881 000 311 174 212 970 000
Asia 2 617 155 14 499 325 6404 7359 1 675 921 10 670 688 1 397 795 1 343 293 1 681 843 1 523 401 000
Caribbean 2156 257 317 13 919 9195 3001 236 602 100 273 89 401 000 897 1 019 000
Europe 688 286 263 487 17 230 18 536 1 185 919 488 408 609 885 690 887 000 139 648 167 956 000
Central 8750 2 278 678 31 657 7240 27 700 1 649 772 230 600 220 369 000 104 363 117 170 000
America
North NA 804 482 NA 21 392 NA 1 720 950 225 709 258 776 000 603 769 513 547 000
America
South 1790 3 419 069 14 441 10 482 2585 3 583 840 332 041 272 811 000 455 509 444 047
America
Oceania 694 596 60 000 13 715 8333 952 645 50 000 15 872 20 145 000 96 692 94 439 000

P, pulses; D, dry beans.

The import export data are for 2011 and the rest for 2012.
Data from FAO (2012 and 2011) Statistical Databases. Rome, Italy: Food and Agricultural Organization of the United Nations (http://faostat.org/site339/default.aspx and http://faostat.
fao.org/site/535/default.aspx#ancor; accessed on 01.05.2014).

Seedcoat (testa)
Cotyledon
Raphe
Plumule
Hilum Hypocotyl
Radicle
Micropyle

(a) (b) Micropyle

(c) Hilum

Outer
integument
Seedcoat

Inner
integument

Epidermis

Cotyledon
Storage
parenchyma
cells
(d)

Figure 1 Dry bean (Phaseolus vulgaris) seed: (a) external side view, (b) external face or edge view (viewing at hilum side), (c) cross section (one
cotyledon removed), and (d) detailed cross section across seed coat and cotyledon. Reproduced from Encyclopedia of Food Sciences and Nutrition,
2nd edn. (2003), p. 407, Elsevier Ltd.

parts of legume seeds are shown in Figure 1. The outermost
layer of the seed is the seed coat or testa. The external seed
structure includes the hilum, micropyle, and raphe. The hilum
is a scar-like structure (usually oval shaped) near the middle
edge where the seed breaks away from the stalk. The micropyle
is the small opening in the seed coat where originally the
pollen tube enters the valve. The raphe is the ridge at the side
of hilum opposite to the micropyle and represents the base of
the stalk that fuses with the seed coat upon seed maturation. In
most legumes, the endosperm is short-lived and shrinks to a
300 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview

thin layer surrounding the cotyledons (or embryo). On soak-
ing the seeds, the endosperm is easily removed along with the
seed coat. The remainder (embryo) of the seed consists of
shoot (which contains two cotyledons) and a short axis
above and below the cotyledons and terminates in the shoot
tip. The plumule or embryonic stem is well developed in the
resting seed and lies between two cotyledons.
The outermost layer of seed coat is the cuticle, which has
papillae or papillae-like growth in some legumes (e.g., green
gram), but in most legumes, it is a smooth structure. The
thickness of the seed coat is quite variable depending upon
the type of bean. Generally, seeds containing thick seed coats
tend to have high fat content. Both the hilum and micropyle
are important in water imbibition by the testa. Palisade cells
derived from the outer epidermis of the outer integument,
which are next to the cuticle, are either loosely packed or
densely packed, depending upon the seed maturity, and may
affect seed hydration. Next to palisade cells are the hourglass
cells. The shape of hourglass cells can vary from bottle-shaped
(guar) to dumbbell-shaped (broad beans) to hourglass-
shaped (soybeans). Only a few legume species (such as
Dolichos, Cajanus, and Vigna) have more than one layer of
hourglass cells. The remainder of the testa contains primarily
mesophyll cells.
Legume cotyledons are primarily composed of parenchyma
cells, which have variable size (70100 mM) and act as storage
sites for most nutrients. Each cell of the cotyledon is bound by
the cell wall and the middle lamella. Vascular bundles in cotyle-
dons generally are devoid of any filling material. Vascular bundles
are often used as key structures in the identification of different
plant types.
Chemical and Nutritional Composition
The majority of nutrients in dry beans are primarily located in
the cotyledons and account for up to 90% of the total nutritive
value. Typically, dry beans provide 12551464 kilojoules (kJ)
per 100 g of dry seeds. The main constituents of cotyledons are
proteins and carbohydrates and, respectively, account for
1525% and 5075% of the total seed weight. The remaining
components consist of lipids, minerals, fiber, and vitamins.
With the exception of oilseeds, dry beans generally contain
low amounts (13% of seed weight) of lipids. Although most
minerals are present in cotyledons, some (such as calcium and
iron) may be present in the seed coat in significant proportion.
Typical nutrient composition of several Phaseolus beans is
shown in Table 3. Dry beans contain not only significant
amounts of nutrients but also several undesirable components
and attributes: inhibitors of enzymes such as trypsin, chymo-
trypsin, subtilisin, amylases, and elastase; lectins; phenolic
compounds including tannins; phytates; toxic amino acids
mimosine and djenkolic acid; cyanogenic glycosides, which
produce HCN; flatulence-causing oligosaccharides raffinose,
stachyose, and verbascose; lipoxygenases, which catalyze the
development of rancidity; and off odors often described as
beany, grassy, painty, cardboard-like, and chalky.

Table 3 Proximate composition of Phaseolus beans

Bean Moisture (%) Protein (%) Carbohydrates (%) Fat (%) Ash (%) Crude fiber (%)

Adzuki 11.00 20.20 49.80 1.90 4.39 4.90

Black beauty 10.41 22.87 70.79 4.48 1.86
Black gram 10.210.9 19.724.0 56.663.4 1.31.6 3.23.4 4.46.4
California small white 9.65 25.90 58.00 0.25
Cranberry 12.71 23.43 71.26 1.09 4.22
Great Northern Lima 8.513.3 21.024.37 61.271.07 1.03.48 3.54.86 6.70
Baby 13.3020.40 20.40 62.10 0.80 3.40 6.00
Large 8.90 22.30 63.80 0.80 4.20 7.40
Mung
Green 17.92 27.12 62.85 1.53 4.01 4.50
Black 13.64 25.68 64.20 0.45 4.32 5.30
Navy 9.418.2 23.1324.65 61.266.19 1.54.3 2.904.27 3.46.6
Pinto 9.0514.70 18.824.97 61.869.47 1.23.6 3.074.10 3.96.3
Red kidney
Light 10.52 20.89 73.20 1.52 4.39
Dark 13.22 20.32 73.68 1.58 4.42
Rice bean 18.025.0 60.077.0 1.01.6 3.84.3 3.34.8
Roshina G2 9.8911.11 25.7726.30 63.3364.02 1.852.00 3.193.79 4.65.1
Roshina pink 4.90 19.40 68.80 3.50 3.40 4.60
Sanilac 11.61 18.98 75.09 1.65 4.28
Tepary 21.025.0 70.073.0 0.80.9 4.14.8
Small red 13.12 22.45 71.97 1.43 4.15
Small white 13.03 19.73 74.34 1.99 3.94
Viva pink 12.69 21.30 73.23 1.06 4.41

Data expressed on dry weight basis.

Data compiled from Sathe SK, Deshpande SS, and Salunkhe DK (1984) Dry beans of Phaseolus: A review. Part 1. Chemical composition: Proteins. CRC Critical Reviews in Food
Science and Nutrition 20: 146; Salunkhe DK and Kadam SS (1989) Handbook of World Food Legumes: Nutritional Chemistry, Processing Technology, and Utilization, Vols. I
(310 pp.), II (294 pp.), and III (323 pp.). Boca Raton, FL: CRC Press.
 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview 301

Proteins from pH 5.6 to 5.8, depending on phaseolin type. Among

Phaseolus beans, three distinct types of phaseolins named
Phaseolus beans not only are important for their caloric contri-
after cultivars Tendergreen (T), Sanilac (S), and Contender
bution to human diet but also are especially valued for their
(C) have been identified. Screening of 107 cultivars has
protein content since they are a major protein contributor to
revealed that S-, T-, and C-type phaseolins accounted for
the human diet on a global basis. In certain parts of the world,
69%, 25%, and 6%, respectively, of the total cultivars. These
they are the sole source of dietary protein. Dry bean proteins
types can be easily distinguished by one-dimensional or two-
can be classified as storage and metabolic proteins. The origi-
dimensional gel electrophoresis using sodium dodecyl sulfate
nal protein classification proposed by Osborne was based on
(SDS) polyacrylamide gel electrophoresis (PAGE) for one-
solubility of proteins in a series of solvents. In this scheme, the
dimensional or isoelectric focussing in first dimension
water- and dilute salt-soluble proteins were termed as albu-
followed by SDS-PAGE in the second dimension for two-
mins and globulins, respectively. Dry beans contain 4060%
dimensional gel electrophoresis. The C-type phaseolin is
globulins and 2040% albumins based on Osbornes protein
believed to have originated from T and S types. Regardless of
classification. Globulins are exclusively storage proteins, while
the type of phaseolin, phaseolin contains 35% carbohydrates
albumin fraction contains both storage and metabolic pro-
and its amino acid composition is dominated by acidic amino
teins. The protein content of dry beans is usually calculated
acids (3040% of total). Typically, the N-glycosylation sites in
by multiplying Kjeldahl nitrogen content by a factor of 6.25.
phaseolin polypeptides occur at amino acid residue numbers
Because dry beans contain 1015% of total nitrogen as non-
252 and 341. Sulfur-containing amino acids (notably methio-
protein nitrogen, most dry bean protein values are typically
nine) are the limiting amino acids in phaseolin as well as other
overestimated by 12%.
major storage proteins in Phaseolus beans. The secondary struc-
The major storage proteins in dry beans have been identi-
ture of phaseolin in 0.5 M NaCl typically has a low a-helix
fied by several names and therefore their nomenclature is
content (10%) and b-turns (9.0%) and large amount of b-sheet
somewhat unclear. Based on the nomenclature using ultracen-
(48.0%) and random coils (33.0%). Native phaseolin is quite
trifugation sedimentation coefficient (S), dry beans contain
resistant to digestive proteases such as pepsin, trypsin, and
both 7S (vicilin-like) and 11S (legumin-like) storage proteins.
chymotrypsin and is degraded to polypeptides with MWs
Depending on the bean variety, the relative proportion of these
24 00028 000. However, heat-denatured phaseolin is easily
two types of proteins varies considerably. The 11S-type pro-
digested by these proteases.
teins typically are nonglycosylated proteins with estimated
molecular weight (MW) in the range 300 000400 000. They
Lectin
are usually composed of six subunits (MW 60 000), each con-
Bean lectins agglutinate erythrocytes due their ability to bind
sisting one acidic (MW 40 000) and one basic (MW 20 000)
with cell surface glycoproteins and glycolipids. Although the
polypeptide linked by disulfide bond(s). Usually, 11S proteins
precise function of lectins in beans is not known, they are
are present in minor amounts in Phaseolus beans. The 7S glob-
thought to offer protection to the plant. Since lectins are
ulin in Phaseolus beans has also been referred to as glycoprotein
toxic, they are of nutritional concern. Lectins occur in both
II, globulin 1, euphaseolin, globulin, and phaseolin. Depend-
albumin and globulin fractions. Certain bean cultivars lack
ing on the type of bean, the 7S globulin type and quantity vary
lectins. When present, lectins represent 612% of total protein.
considerably. In Phaseolus beans, the 7S globulins are, how-
In addition to agglutinating activity, many bean lectins have
ever, the major storage proteins and account for 4060% of the
mitogenic activity. Most lectins of Phaseolus beans have sub-
total proteins. The three major types of 7S proteins that have
units of MWs 29 00036 500 with isoelectric pH in the range
been identified, biochemically purified, and characterized are
4.97.9 (most are in the pH range 56). The majority of the
(1) phaseolin, (2) lectin (also called glycoprotein II, phytoag-
native lectins have tetrameric nature (MWs 100 000150 000)
glutinins or phytohemagglutinins, and protein II), and (3)
although some (e.g., lima beans) have dimeric nature. The
arcelin in wild bean accessions from Mexico (named after
majority of Phaseolus lectins have 46% carbohydrate, low
town Arcelia in Mexico, where some of the accessions were
sulfur-containing amino acids, and sugar specificity toward
collected). All the 7S globulins are glycosylated and contain
D-acetylgalactosamine. Lectins are very resistant to common
D-mannose and D-glucosamine as the major sugar constituents.
digestive proteases and are slowly hydrolyzed in vitro even
after extensive heat denaturation. Proper moist-heat denatur-
Phaseolin ation can completely inactivate the biological activity of lectins
Phaseolin is the major globulin in domesticated Phaseolus and therefore can render them nontoxic.
beans. It is a trimeric, vicilin-like 7S globulin known to exhibit
polymorphism. The polypeptide polymorphism is believed to Arcelin
be not only due to the differential glycosylation but also Arcelin was first discovered in wild accessions of Mexican
because phaseolin polypeptides are encoded by a small multi- Phaseolus beans. It also occurs in lines that contain phaseolin
gene family. Phaseolin is soluble in 0.5 M NaCl at all pH as well as lectin. Because it is present in equal or greater levels
values. It undergoes reversible pH-dependent dissociationas- than phaseolin in certain lines, it is one of the major storage
sociation with sedimentation coefficients of 3.0S (pH 12.0), proteins in Phaseolus beans. The MWs of arcelin subunit poly-
7.1S (pH 7.0), and 18.2S (pH 3.6) known as peptides (MW peptides range from 35 000 to 42 000 depending on the
44 000), protomers (MW 163 000), and tetramers of proto- variant and are more basic than both lectin and phaseolin.
mers (MW 653 000). Phaseolin consists of a group of subunit The native arcelin has an MW of 89 000 and is therefore a
polypeptides with MWs 43 00054 000 and isoelectric points dimeric protein. Arcelin has many similarities with lectin
302 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview

(including agglutinating activity) with respect to chemical shown to be 50 000. Appropriate moist-heat treatment (such
composition. as cooking or autoclaving) can inactivate both the protease and
amylase inhibitors.

Other proteins
Phaseolus beans contain trypsin inhibitors (many of them also
inhibit chymotrypsin), amylase inhibitors, lipoxygenases, and
several other minor protein components. Most of these pro-
teins are a part of albumins. Trypsin and chymotrypsin inhib-
itors in Phaseolus beans typically account for up to 10% of the
total proteins and are generally rich in sulfur amino acids. The
MWs of these inhibitors range from 2000 to 23 000. Most
Phaseolus beans lack Kunitz-type (inhibitors with 170200
amino acids with MW of
20 000) trypsin inhibitors. Amylase
inhibitors in dry beans have been characterized from only a
few cultivars and therefore not yet extensively studied. The MW
of kidney bean amylase inhibitor (a glycoprotein) has been
Allergenic Proteins
With heightened concerns about food allergies and the
increased frequency of food recalls due to actual or suspected
presence of allergen in food, food-induced allergens are of
increased interest to consumers and regulatory agencies alike.
Since any food protein could be a potential allergen, it is
important to recognize that dry bean proteins may be aller-
genic. Compared to aeroallergens, such as pollen, perhaps with
the exception of peanut and soybean proteins, very little is
known about dry bean protein allergens. A few dry bean pro-
teins have been identified to be allergens (Table 4) and even

Table 4 Recognized legume allergens

MW (kDa) Allergen
Legume SDS-PAGE Identity designation Seq.a Accession number PubMed index

Dry beans
Chickpea
Cicer 10106 11527247,
arietinum 10705224
Cowpea
Vigna 41, 55 Albumins 12546052
sinensis
Grass pea
Lathyrus 21, 28, 46 11295670
sativus
Green pea
Pisum 20, 2030, Vicilin-like proteins 12589366
sativum 33, 50
1.8 Glycoprotein 991447
French 8.89 Nonspecific lipid transfer protein type 1 C GQ434007.1
(green) (ns-LTP-1)
bean Nonspecific lipid transfer protein 1b C GQ434008.1
precursor
Lentil
Lens 1284 11080720,
culinaris 10604559,
9893199
48 Vicilin-like protein Len c 1 C
66 Seed-specific biotinylated protein Len c 2 P 11080720
Mung bean 16 Pathogenesis-related protein, PR-10 C AY792956.1
Vigna radiata Bet v 1 family member
53 8S globulin (vicilin) C Q198W3, B1NPN8
30 Albumin C Q43680
18 Cytokinin-specific binding protein (CSBP), C AB012218
Bet v 1 family member
Oil seeds
Castor bean
Ricinus 11.2 2S albumin Ric c 1 C P01089 9430499
communis
12 2S albumin Ric c 3 P 9430499
11S crystalloid protein Ric c 2 3392372
47/51 Protein doublet 3392372
Lupin
Lupinus 30175 10359910
albus
Peanut
 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview 303

Table 4 (Continued)

MW (kDa) Allergen
Legume SDS-PAGE Identity designation Seq.a Accession number PubMed index

Arachis 63.5 Vicilin-like protein Ara h 1 C L34402, L38853, 7560062

hypogaea AF432231
17 Conglutin Ara h 2 C L77197, AY158467, 14538941,
AY117434 11295663
60 Glycinin Ara h 3 C AF093541, 10021462
AF510854
37 Glycinin Ara h 4 C AF086821, 10474031
AF510854
15 Profilin Ara h 5 C AF059616 10474031
15 Hom: conglutin Ara h 6 C AF092846 10474031
15 Hom: conglutin Ara h 7 C AF091737 10474031
17 Pathogenesis-related protein (PR-10) Ara h 8 C AY328088
30, 50, 68 Oleosin oligomers 12144563
12 Oleosin monomer 12144563
Soybean
Glycine max 7 Hydrophobic seed protein (HPS) Gly m 1A/1B P AAB34755, 7603155
AAB34756
8 Hull protein Gly m 2 P A57106
14 Profilin 1, Profilin 2 Gly m 3 C AJ223981, 10589015
AJ223982
17 (SAM 22) PR-10 prot. Gly m 4 C X60043 12417891
21 Trypsin inhibitor 8836337, 11799388
26 Vicilin-like glycoprotein Gly m Bd C AB046874 11267676
28k
34 Oil body associated glycoprotein Gly m Bd C AB013289 11227798
30k
Alpha-subunit of beta-conglycinin Gly m Bd 7787297
60k
22 G2 glycinin 11112856
15 Glycinin G1 acidic chain 11146387
7, 12, 20, 39, Soy lecithins 11752879
57
12 Methionine-rich protein (MRP), 2S P 11752879
albumin
20 Soybean Kunitz-type trypsin inhibitor P 11752879
(SKTI)
a
Sequence source: P, protein; C, cDNA.
Compiled in part from the International Union of Immunological Societies, Allergen Nomenclature Sub-committee http://www.allergen.org/list.htm.

fewer have been characterized at molecular level. Phaseolin,

the major storage globulin in dry beans, has not yet been
reported to be an allergen. Among the legume proteins identi-
fied to be allergens, majority appear to be storage proteins such
as 2S albumins (e.g., Ric c 1, Ric c 3, and soybean MRP), 7S or
vicilin-like proteins (e.g., Ara h 1, b-conglycinin, and Len c 1),
and 11S or legumin-like proteins (e.g., soybean glycinin, Ara h
3, and Ara h 4), some of which are glycosylated (Ara h 1 and
Gly m Bd 28 k), while others are not (soybean glycinin). Since
typical thermal food processing does not significantly hydro-
lyze food proteins, linear stretches of amino acids responsible
for IgE binding, the linear epitopes, are expected to survive
normal food processing operations. For these reasons, food
processing is often unable to completely inactivate food aller-
gens. It is therefore also not surprising that some investigators
find many legume allergens to be acid- and heat-resistant.
Carbohydrates
Total carbohydrates in Phaseolus beans contribute 5070% of
the seed weight and include mono-, di-, and oligosaccharides;
starch; and other polysaccharides. Starch is the most abundant
nutrient in Phaseolus beans accounting for up to 7080% of
total carbohydrates. Among the simple sugars, oligosaccha-
rides (raffinose, stachyose, verbascose, and ajugose) are the
major constituents (up to 10% of seed weight) and are at
least partially responsible for flatulence production. Crude
fiber is primarily composed of cellulose, hemicellulose, lignins
(not a carbohydrate), and other nonstarchy polysaccharides
such as arabinogalactans, arabinoxylans, glucomannans, galac-
tomannans, and pectins. The hypocholesterolemic effect of dry
beans is partially attributed to the presence of nonstarchy poly-
saccharides. Phaseolus bean starch granules are quite variable in
304 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview
shape (round, oval, oblong, elliptical, spherical, kidney-shape-
d, and irregular) and size (560 mm) and typically contain
1045% (of total starch) amylose. The average degree of poly-
merization for amyloses and chain lengths for amylopectins of
Phaseolus bean starches ranges from 1600 to 1900 and 22 to 26,
respectively. Based on x-ray diffraction spectra, Phaseolus dry
bean starches are mostly C type (mixture of A (typical of cereal
starches) and B (typical of root and tuber and high-amylose
cereal starches) types). These starches have restricted swelling,
gelatinization temperature range of 6089  C, high solution
viscosities, and good thermal stabilities. Upon gelatinization,
they produce opaque gels. Dry bean starches (especially if
cooked) are well digested (digestibility is comparable to those
of many cereal and tuber starches) by humans. However, as dry
bean starches are digested slowly, they are hypoglycemic and
therefore useful in the diets of diabetics. The nonstarchy poly-
saccharides are not digestible in many mammals due to the
lack of required enzymes in their digestive tracts and therefore
contribute to the total dietary fiber. These nonstarchy poly-
saccharides help increase viscosity and may thereby lower
digestion rate of starch (i.e., lower glycemic index). Further,
they may assist removal of dietary lipids (i.e., hypocholester-
olemic effect).
Vitamins and Minerals
Phaseolus beans are a good source of B vitamins, especially
thiamin, riboflavin, niacin, and folacin. Typically, thiamine,
riboflavin, niacin, and folacin contents (on a dry weight basis)
of Phaseolus beans are, respectively, 0.51.14, 0.10.25,
0.43.14, and 0.0370.676 mg per 100 g. Vitamin E content
ranges from 0.72 to 1.97 mg per 100 g and B6 content ranges
from 0.2 to 0.659 mg per 100 g. Phaseolus beans are not good
sources of vitamin A and C. Phaseolus beans are excellent
sources of several minerals including Ca, Fe, Cu, Zn, P, K,
and Mg. Typically, raw beans contain 70260, 0.51.40,
3.3413.5, 160320, 1.02.1, 380570, 13201780, in vivo model studies to decipher possible beneficial effect(s).
4.021.0, and 1.96.5 (all expressed as mg per 100 g, dry
weight basis) of Ca, Cu, Fe, Mg, Mn, P, K, Na, and Zn, respec-
tively. The low sodium and high potassium content of raw
beans makes them a desirable constituent of human food
(especially for people with hypertension).
Antinutritional Factors
In addition to protease inhibitors and amylase inhibitors, Pha-
seolus beans contain several other antinutritional and
antiphysiological factors such as phytic acid, tannins, cyano-
genic glycosides, saponins, and allergens. Because these com-
ponents are usually present in small quantities (less than 5% of
the total seed weight), they do not pose a serious health hazard
under normal conditions (i.e., when beans are a part of total
diet and are properly processed prior to consumption). Of
these antinutritional factors, phytates and tannins are of par-
ticular concern because both of them are heat-stable and can-
not be easily removed from beans during normal home
processing. Phytate is a general term used for mono- to dodeca
anions of phytate along with esters lower than hexaphosphate.
CaMg salts of phytic acid (myoinositol 1, 2, 3, 4, 5, 6-hexakis
(dihydrogen phosphate)) are referred to as phytin. In most dry
beans, phytate phosphorus accounts for up to 80% of the total
phosphorus. The amount of phytate in Phaseolus beans ranges
from 0.6% to 2.1% (by weight) of the total seed weight.
Because phytates are chelating agents, they may interfere in
mineral utilization. Germination, fermentation, and soaking
followed by cooking (if both soak and cook waters are dis-
carded) are effective methods of removing phytates (5080%
reduction). Tannins (especially condensed tannins) are heat-
stable compounds and are present in Phaseolus seeds (espe-
cially in colored varieties) up to 2% of total seed weight.
Because of their ionic character, they may interact with other
constituents (notably proteins) and adversely affect the nutri-
tional bioavailability of that constituent. Tannins are thought
to offer protection to the plant from insects and pests.
Lipids
Phaseolus beans contain 13% lipids (by weight) depending
upon the species. Neutral lipids (3050% of total) and phos-
pholipids (2535% of total) are the major constituents, and
glycolipids may account for up to 10% of the total lipids.
Regardless of the variety, Phaseolus bean lipids primarily con-
tain palmitic, oleic, linoleic, and linolenic acids. Polyunsatu-
rated fatty acids and saturated fatty acids typically account for
5587% and 1228% of total lipids.
Beneficial Bioactive Compounds
Continuing research on raffinose oligosaccharides, phytates,
tannins, simple phenolic compounds, sterols, protease inhib-
itors, and several other major and minor constituents of dry
beans suggests that some of these compounds, under appro-
priate conditions, may offer beneficial effects for human
health. Numerous reports suggesting the beneficial effects of
one or more components (either alone or in combination)
have appeared. These investigations have used in vitro and
The primary goals of these investigations are to (a) derive
nutritionally beneficial effect for long-term good physical and
mental health and (b) treat/cure disease. Experimental dem-
onstration of causeeffect relationship can be tenuous and
difficult to prove/disprove and may depend on the experimen-
tal design. With the simultaneous presence of several com-
pounds in the same food source, establishing unequivocal
relationship between the compound(s) and their physiological
effect(s) is, at best, very difficult. In recent years, attempts to
investigate the underlying molecular and mechanistic basis for
the reported beneficial effect(s) of tested compound(s)/com-
ponent(s) have therefore often resulted in outcomes with very
limited success. Long-term detailed and fundamental research
is needed to fully realize the potential health benefits of dry
bean consumption.
Grading, Handling, and Storage
Dry beans are usually harvested at maturity. The seeds are
removed from the pods either manually or mechanically and
cleaned to remove dirt, stalks, leaves, blemished, wrinkled, and
broken seeds and packaged prior to storage. The grading of

seeds is usually based on external characteristics such as color,
gloss, seed size, seed soundness, seed firmness, and presence of
contaminating substances. The seeds are stored at farmer,
trader, or government levels. Typically, farmers hold up to
8% of harvested seeds until next season so that they can be
used as planting seeds if crop failures occur. In developed and
developing nations, a majority of the seeds are stored by
traders and/or governments to protect against subsequent
crop failures (or low yields), price fluctuations, and fluctua-
tions in supply (change in demand, shortages, and famines).
Losses in seeds occur both pre- and postharvest. The normal
preharvest losses are mainly due to birds and mammals feed-
ing on bean plant seeds. Drought/floods, insects, and rodents
can also contribute to preharvest losses. In developed nations,
preharvest losses are usually small (as low as 1% of the crop). A
majority of the losses occur during postharvest handling and
storage and can range from 8% to as high as 50% of total crop.
It is estimated that as much as 48% of food produced in the
world is lost (due to pre- and postharvest losses). Factors that
influence postharvest losses of legumes include moisture, tem-
perature, respiration rate, insect damage, microbial spoilage,
and damage caused by mites and rodents. Properly packaged
dry beans should be stored at low relative humidity and tem-
perature conditions. High relative humidity and temperature
favor the hard-to-cook beans. These conditions also favor the
growth of molds and insects. Three major insect genera that
cause much of the damage to stored legumes are Bruchus,
Acanthoscelides, and Callosobruchus. Dehusked, split stored
pulses are also damaged by Rhyzopertha, Trogoderma, and Tri-
bolium species. Usually, pests seem to have preference for the
type of bean they infest although the basis for such preference
(or the lack of it) has not been elucidated yet. The major
microbial problem during bean storage is contamination by
aflatoxin-producing molds (Aspergillus flavus and Aspergillus
parasiticus). Mites can consume food up to their own weight
(68 mg) and because of their large numbers can cause serious
losses. Rodents cause twofold damage to stored legumes by not
only consuming but also contaminating (up to 20 times the
amount they would eat). Because rodents are carriers of many
communicable diseases, they pose serious damage to stored
beans. The species that most commonly cause damage include
Rattus rattus, R. norvegicus, Bandicota indica, B. bengalensis, and
Mus musculus.
Processing and Food Uses
Phaseolus beans are processed and used in a variety of ways. The
processing of beans is mostly at the household level in devel-
oping and underdeveloped countries, while in most developed
countries, the majority of the processing is done at the indus-
trial level. Home processing methods include milling, soaking,
cooking, frying, germination, fermentation (either alone or in
combination with cereals), roasting, puffing, parching, extru-
sion and frying, and toasting. The method(s) used for home
processing depends on regional preference for bean variety and
the desired end product. For example, mung beans in sprouted
form are popular on global scale and therefore germination is
one of the preferred household processing methods used.
Black gram, on the other hand, is extensively used for the
THE LEGUMES AND PSEUDOCEREALS | Beans: Overview 305
preparation of idli, a breakfast food popular in India and Sri
Lanka, after fermenting it with rice. Industrial processing
includes freezing for such beans as green French beans and
snap beans, milling (production of flours and high-protein
flours), baking (baked beans), cooking and frying (refried
beans), and canning (alone in saltwater or tomato juice or in
combination with meats such as beef and pork). In developing
countries, dehusking and splitting to produce dal is also done
at industrial scale.
In developing and underdeveloped countries, Phaseolus
beans are used in numerous ways depending on the type of
bean and regional preference. They may be eaten as raw,
immature seeds; cooked as green vegetables (such as French
beans); consumed as part of salads; used in making curry; used
as a soup ingredient; cooked, mashed, and mixed with condi-
ments and spices and used as gruels and porridges; prepared as
pastes to be extruded to prepare fried snack products; sprouted;
puffed or roasted and eaten as snack foods; and fermented to
prepare numerous fermented products.
In developed countries, Phaseolus beans are consumed as
salad and soup ingredient, sprouts, canned, frozen, and refried
beans. They are also extensively used in the preparation of
Mexican-style preparations, such as burrito, chimichanga,
taco, bean dips, and tamale, and often canned with meats
such as beef and pork. In many South American countries,
cooked black beans are a preferred part of breakfast. With
increased ethnic populations in the Western world, increased
emphasis on vegetarian foods, and recommended reduction of
meats and animal foods in the human diet, opportunities need
to be explored to incorporate dry beans in food products for
improved and nutritious food products.
Although there is a good potential for the preparation of
protein concentrates and isolates and the development of food
starches, Phaseolus beans have not been used on large scale for
such purposes. In many countries, especially the developed
ones, Phaseolus beans have been extensively used as animal
feed. In developing and underdeveloped countries, the green
foliage, deseeded pods, and roots and shoots of bean plants are
used as natural fertilizers especially after composting. Because
the legume roots fix nitrogen, they help conserve soil quality.
For this reason, in many developing countries, they are exten-
sively used for soil quality conservation.
Exercises for Revision
What distinguishes cereals from legumes?
What is the difference between the terms legume and
oilseed?
Which type of beans are the most widely consumed in the
world?
Which compound(s) account for 80% phosphorus in dry
beans?
What is the advantage of using dry beans with cereals in
foods?
What is the advantage of the nonstarchy polysaccharides
present in dry beans in human nutrition?
What is the difference between the terms pulse and
legume?
306 THE LEGUMES AND PSEUDOCEREALS | Beans: Overview
Exercises for Readers to Explore the Topic Further
What are the possible beneficial actions of bioactive com-
ponents of dry bean?
What are the factors that result in pre- and postharvest
losses?
How does the ionic character of phytic acid influence its
interaction with dietary minerals and proteins?
In what ways can the dry bean utilization be improved?
See also: Agronomy of Grain Growing: Lentil: Agronomy;
Breeding of Grains: Lentil: Breeding; Food Grains and the
Consumer: Cultural Differences in Processing and Consumption;
Proteins: The Protein Chemistry of Dicotyledonous Grains; The
Basics: Taxonomic Classification of Grain Species; The Legumes
and Pseudocereals: Chickpea: Overview; Grain Legumes and Their
Dietary Impact: Overview; Pea: Overview.
Further Reading
Broughton WJ, Hernandez G, Blair M, Beebe S, Gepts P, and Vanderleyden J (2003)
Beans (Phaseolus spp.) Model food legumes. Plant and Soil 252: 55128.
Derbyshire E, Wright DJ, and Boulter D (1976) Legumin and vicilin, storage proteins of
legume seeds. Photochemistry 15: 324.
Deshpande SS (1992) Food legumes in human nutrition: A personal perspective. CRC
Critical Reviews in Food Science and Nutrition 32: 333363.
Doughty J and Walker A (1982) Legumes in Human Nutrition, p. 152. Rome, Italy: Food
and Agriculture Organization of the United Nations Food and Nutrition Paper 20.
FAO (2013) http://faostat.fao.org/site/567/DesktopDefault.aspx?PageID567#ancor
(accessed 22.02.2014).
FAO (2013) Dimensions of Need Staple Food: What Do People Eat? http://www.fao.
org/docrep/u8480e/u8480e07.htm (accessed 22.02.2014).
Geil PB and Anderson JW (1994) Nutrition and health implications of dry beans: A
review. Journal of the American College of Nutrition 13: 549558.
Guillon F and Champ MM (2002) Carbohydrate fractions of legumes: Uses in
human nutrition and potential for health. British Journal of Nutrition
88: S293S306.
Kami JA, Velasquez VB, Debouck DG, and Gepts P (1995) Identification of presumed
ancestral DNA sequences of phaseolin in Phaseolus vulgaris. Proceedings
of the National Academy of Sciences of the United States of America
92: 11011104.
Muntz K (1998) Deposition of storage proteins. Plant Molecular Biology 38: 7799.
Osborn TC (1988) Genetic control of bean seed proteins. Critical Reviews in Plant
Sciences 7: 93116.
Patwardhan VN (1962) Pulses and beans in human nutrition. American Journal of
Clinical Nutrition 11: 1230.
Reddy NR, Pierson MD, Sathe SK, and Salunkhe DK (1984) Chemical, nutritional and
physiological aspects of dry bean carbohydrates: A review. Food Chemistry
13: 2568.
Reddy NR, Pierson MD, and Salunkhe DK (1986) Legume-Based Fermented Foods,
p. 254. Boca Raton, FL: CRC Press.
Salunkhe DK and Kadam SS (1989) Handbook of World Food Legumes: Nutritional
Chemistry, Processing Technology, and Utilization. Boca Raton, FL: CRC Press
vols. I (310 pp.), II (294 pp.), and III (323 pp.).
Sathe SK (2002) Dry bean protein functionality. Critical Reviews in Biotechnology
22: 175223.
Sathe SK, Deshpande SS, and Salunkhe DK (1984) Dry beans of Phaseolus: A review.
Part 1. Chemical composition: Proteins. CRC Critical Reviews in Food Science and
Nutrition 20: 146.
Siddiq M and Uebersax MA (eds.) (2013) Dry Beans and Pulses: Production,
Processing and Nutrition, pp. 398. Ames, IA: Wiley.
 Buckwheat: Overview
YZ Cai, China National Seed Group, Beijing, China; Huazhong Agricultural University, Wuhan, China
H Corke, The University of Hong Kong, Hong Kong, Pokfulam, China; Hubei University of Technology, Wuhan, China
D Wang, Huazhong Agricultural University, Wuhan, China
WD Li, Shanxi University, Taiyuan, China
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Cultivated buckwheat comprises two species of the genus
Fagopyrum, namely, F. esculentum Moench (common buck-
wheat) and F. tataricum L. (Tartary or bitter buckwheat).
Originating in China and widely produced, mostly in Asia
and in eastern Europe, buckwheat produces a pseudocereal
grain that can be dehulled and milled to flour, for use in a
wide range of food products.
The composition and properties of buckwheat flour,
derived from its components of starch, protein, and various
therapeutic components such as flavonoids and phytos-
terols, give a wide scope for development of healthful
grain-based products.
Learning Objectives
To understand the origin, geographic distribution,
production, and potential of buckwheat as a traditional
grain with emerging uses in healthful products
To understand the composition and properties of the major
components of buckwheat flour (starch and protein) and
how these contribute to the nutritional and physical char-
acteristics of products
To understand the diverse array of therapeutic components
present in buckwheat seed and how these can be manipu-
lated for maximum benefit
Introduction
Buckwheat originated in China and is widely cultivated as a
minor crop in many places of the world. It is a pseudocereal
crop, belonging to the genus Fagopyrum of the family
Polygonaceae. There are two cultivated species, namely, com-
mon buckwheat (Fagopyrum esculentum) and Tartary buck-
wheat (Fagopyrum tataricum). Common buckwheat is
most widely cultivated and utilized, while Tartary buckwheat
is mainly grown and consumed in China. The major
buckwheat-producing and buckwheat-consuming countries
include China, Japan, Korea, Nepal, India, Russia, Ukraine,
Poland, Hungary, Slovenia, Italy, Canada, Brazil, and the
United States. Buckwheat is mostly consumed in the form of
groats and flour, which are used as material for porridge,
noodles, bread, pancakes, spaghetti, and other food items.
Many traditional buckwheat foods are still popular in some
countries, such as Japanese soba and wantuo and helao in
China. Buckwheat is rich in protein of excellent quality and
starch with special properties that are different from other
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00034-6
cereals. In addition, it also contains high levels of fiber, min-
erals, and vitamins and other bioactive compounds (e.g., fla-
vonoids, phytosterols, D-chiro-inositol, myo-inositol, free and
bound phenolic acids, and phenylpropanoid glycosides) with
positive therapeutic effects on the human body. Buckwheat
with both edible and pharmaceutical values has increasingly
attracted attention because of its positive effects on some
chronic disease conditions, such as hypertension, hypercholes-
terolemia, diabetes, and other cardiovascular diseases. Buck-
wheat has further potential for use as an important functional
food source.
General Aspects of Buckwheat Production
Buckwheat grows mainly in the northern hemisphere, notably
in Russia, China, Japan, India, Poland, Ukraine, Canada, and
the United States. Buckwheat is also planted in the southern
hemisphere, for example, in Brazil, Australia, and South Africa.
In 1993, Russia was the biggest producer of buckwheat in the
world, with a planting area of more than 4 million hectares
(Mha), mainly in the eastern and northeastern parts of the
country. It is reported that in 1990, buckwheat yield per hect-
are reached 3400 kg in Russia. China has the longest history
(since the AD seventh century) of buckwheat cultivation and
now ranks second in world production with an average annual
cultivation area of
1.33 Mha and production fluctuating in
the range of 0.50.9 million tonnes (Mt). Average yield per
hectare is 527 kg for common buckwheat and 1294 kg for
Tartary buckwheat in 1986. Major production areas of buck-
wheat in China include four production regions, distributed in
Inner Mongolia, Shanxi, Gansu, Shaanxi, Sichuan, Yunnan,
and Guizhou. From 2006 to 2010, the harvested area and
total production of buckwheat in China and the world
decreased by year (Table 1). However, in recent years, the
planting area and total production of buckwheat in China
have increased, close to 1 Mha and around 0.9 Mt per year.
The total production of buckwheat in China has accounted for
about 40% in the world. Moreover, China is the biggest pro-
ducer of Tartary buckwheat in the world, with over 90%
production.
Biological Aspects of Buckwheat
Buckwheat, a dicotyledonous plant, belongs to the genus Fago-
pyrum of the family Polygonaceae. Fagopyrum has 15 species,
including two groups: (1) annual species such as F. esculentum
Moench (common buckwheat), F. tataricum L. (Tartary or bit-
ter buckwheat), and Fagopyrum giganteum Krotov and (2)
307
308 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview

perennial species such as Fagopyrum cymosum Meissn and Fago-
pyrum suffruticosum Fr. Schmidt. Among these species, only
F. esculentum and F. tataricum are cultivated species while
others are wild species. Buckwheat has broad ecological adapt-
ability and can grow well in adverse soil and climatic condi-
tions; in particular, Tartary buckwheat can grow in poor soils
even at low temperature and low precipitation, such as in high-
altitude mountainous regions.
Common buckwheat is characterized by petiolate leaves,
with terminal and axillary flowers, and corymbose or paniculate
cyme. Tartary buckwheat has narrow-shaped leaves, more
branches, and smaller flowers than common buckwheat, with
inconspicuous greenish-white sepals. Buckwheat flowers are
indefinite inflorescences that are white, pink, or yellow. Because
of its flourishing flowers, buckwheat is cultivated in China to
support the production of honey, considering one of the highest-
yielding honey plants. Buckwheat grain is actually a fruit
(achene) whose color ranges from a glossy or dull gray to
brown or black. Buckwheat grain (achene) consists of hull (peri-
carp) and kernel or seed (testa, aleurone layer, embryo, and
endosperm). Such achenes are mostly triangular,
49 mm
long with 1000 grain weight ranging from 15 to 35 g. Common
buckwheat grain is generally bigger and huskier than Tartary
buckwheat grain. Common buckwheat has a softer hull than
Tartary buckwheat. Common buckwheat grain tastes harsh, and
Tartary buckwheat tastes slightly bitter.
Chemical Composition of Buckwheat
Chemical components of buckwheat include protein/amino
acids, starch, lipids, fiber, minerals, vitamins, and other func-
tional components (e.g., flavonoids, phytosterols, fagopyrins,
D-chiro-inositol, myo-inositol, and phenylpropanoid
glycosides). These are distributed in different parts of the buck-
wheat grain, for example, protein mainly occurs in the aleu-
rone layer and embryo, starch in the endosperm, and fiber, ash,
and flavonoids are normally in the testa and pericarp. Milling
procedures, fineness, and flour yield influence chemical com-
position in the final products of buckwheat milling (Table 2).
Different buckwheat varieties (genotypes) and plant condi-
tions (soil and climate) also cause compositional differences.
Protein and Amino Acids
Buckwheat grain contains not only a high level of protein but
also a good quality or nutritional balance of protein. Protein
content of buckwheat grain and flour is similar to or slightly
higher than that in wheat grain and flour (Table 2) but
significantly higher than those in many other cereals (e.g.,
rice, maize, and sorghum). Amino acid composition of buck-
wheat protein is nutritionally well balanced and is rich in
essential types (e.g., lysine and arginine), compared with
most cereals. Table 3 shows that all amino acids are higher
in buckwheat grain than in wheat grain, except for glutamic
acid and proline.
Carbohydrate
Carbohydrate (mainly starch) is the major component
(5974%) of buckwheat grain (Table 2). Amylose content in
starch generally ranges from 20% to 26%. Starch granules are
215 mm in diameter, round, oval, or polygonal in shape with
a few holes and pits on the surface. Buckwheat starch exhibits a
typical A-type x-ray diffraction pattern and the crystallinity
varies from 38.3% to 51.3%. Buckwheat grains also contain
0.650.76% reducing sugars, 0.791.16% oligosaccharides,
and 0.10.2% nonstarchy polysaccharides.

Table 1 The harvested area and total production of buckwheat in Lipids

China and the world (200610)
Buckwheat grain contains 2.02.6% oil (Table 2). Major fatty
2006 2007 2008 2009 2010 acids are palmitic (16:0), oleic (18:1), and linoleic (18:2)
acids, with average values of 14.8%, 36.5%, and 35.5%, respec-
The harvested area (Mha) tively. Unsaturated fatty acids constitute
7580% of total
China 0.868 0.750 0.725 0.724 0.700 fatty acids.
World 2.740 2.727 2.422 2.016 1.882
Total production (Mt)
China 0.500 0.650 0.600 0.570 0.590 Fiber
World 1.993 2.369 2.181 1.794 1.518
Buckwheat grain contains 9.310.9% crude fiber (Table 2),
Source: Data from FAOSTAT database.
2030% of which is soluble dietary fiber. Common

Table 2 Proximate composition of buckwheat compared with wheata

Composition Buckwheat seeds Buckwheat groats Buckwheat crude flour Buckwheat fine flour Wheat grain Bread wheat flour

Carbohydrate 58.573.5 70.9 67.4 70.7 66.9 74.7

Protein 1014.5 9.7 12.0 10.9 14.0 11.8
Fat 2.02.6 1.8 2.3 1.7 2.1 1.1
Fiber 9.310.9 3.7 7.2 2.6 0.3
Ash 2.02.5 1.7 2.1 1.6 1.9 0.4
Moisture 1214.0 12.3 10.3 13.1 12.5 11.7
a
Percentage on dry weight basis: data from Lin RF (1994) Buckwheat in China (in Chinese). Beijing: Chinese Agriculture Publishing House; Li SQ and Zhang QH (2001) Advances in
the development of functional foods from buckwheat. Critical Reviews in Food Science and Nutrition 41: 451464.
 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview 309

buckwheat grain generally has higher crude fiber than Tartary
buckwheat. Content of crude fiber in buckwheat flour com-
monly depends on the milling process because fiber is mainly
distributed in the testa and pericarp. Crude flour contains more
fiber (3.46.9%) than fine flour (0.11.6%).
Vitamins
Buckwheat grain contains higher levels of vitamin B1
(thiamine), B2 (riboflavin), E (tocopherol), and PP (nicotinic
acid/amide) than most cereals (Table 4). Generally, Tartary
buckwheat has more vitamin B1, B2, and PP, but less vitamin
E than common buckwheat.

Minerals
Buckwheat grain is rich in minerals including Mg, Se, Fe, K, Ca,
Cu, Mn, and Zn. Some studies show that buckwheat flour
contains more Mg (1113 times), Sr (536 times), Li (57
times), Fe (34 times), and K (2.93.6 times) than wheat
flour. Additionally, a high level of Se is detected in Tartary
buckwheat flour (0.43 mg kg1).
Therapeutic Components
Buckwheat is also rich in components with therapeutic effects,
including flavonoids, phytosterols, fagopyrins, D-chiro-
inositol, myo-inositol, phenylpropanoid glycosides, free and
bound phenolic acids, and proanthocyanidins.

Table 3 Amino acid contents of grains from 1505 buckwheat

genotypes compared to wheat grainsa Flavonoids
Common buckwheat Tartary buckwheat Several flavonoids have been identified in Tartary and com-
mon buckwheat, that is, rutin, quercetin, kaempferol, orientin/
Amino acid Range Mean Range Mean Wheat
isoorientin, and vitexin/isovitexin. Rutin is the major and most
Ala 0.280.75 0.51 0.260.79 0.50 0.33 important flavonoid component in buckwheat, which is not
Argb 0.691.72 1.11 0.111.61 1.09 0.51 found in cereals. All parts of the buckwheat plant (seeds, hulls,
Asp 0.102.34 1.09 0.601.84 1.06 0.43 leaves, stems, flowers, and even roots) contain flavonoids. The
Cys 0.030.33 0.17 0.030.44 0.16 0.11 flavonoid components and content in buckwheat differ among
Glu 0.673.49 2.21 0.603.12 2.05 2.44 growth periods and among species and genotypes. Generally,
Gly 0.440.95 0.65 0.072.54 0.64 0.33 the flavonoids in Tartary buckwheat are significantly higher
Hisb 0.170.40 0.26 0.030.50 0.27 0.20
than those in common buckwheat. It is reported that common
Ileb 0.041.65 0.46 0.050.71 0.46 0.27
buckwheat grain contains 413 mg g1 flavonoids, whereas
Leub 0.491.83 0.77 0.081.07 0.75 0.50
Lysb 0.061.96 0.66 0.121.86 0.64 0.29 Tartary buckwheat grain has
40 mg g1. In Tartary buck-
Metb 0.020.33 0.17 0.030.34 0.17 0.16 wheat flowers, leaves, and stems, total flavonoid content
Pheb 0.311.79 0.57 0.050.99 0.55 0.36 reaches over 100 mg g1.
Pro 0.201.06 0.44 0.111.22 0.41 0.92
Ser 0.350.91 0.57 0.050.90 0.58 0.36
Thrb 0.160.67 0.44 0.240.62 0.43 0.21
Phytosterols
Trpb 0.060.18 0.12 0.060.18 0.13
Tyr 0.090.55 0.32 0.070.55 0.33 0.23 Buckwheat contains several kinds of phytosterols, for example,
Valb 0.381.24 0.58 0.261.18 0.58 0.36 b-sitosterol, campesterol, stigmasterol, and isofucosterol.
a b-Sitosterol is the major sterol component, accounting for
Percentage on dry weight basis: data from Lin RF, Zhou MD, Tao YR, Li JY, and Zhang
ZW (eds.) (1992) Proceedings of the 5th International Symposium on Buckwheat,
70% of total sterol content. Some studies report that
2026 August, Taiyuan, China. Beijing: Agricultural Publishing House; Wei YM (1995) dehulled buckwheat groats after lipid extraction have
Quality and Processing of Buckwheat Grain (in Chinese). Xian, China: World 667753 mg kg1 b-sitosterol, 8997 mg kg1 campesterol,
Publishing Corporation. and a trace of stigmasterol. Phytosterols also occur in buck-
b
Essential amino acids. wheat pollen.

Table 4 Comparison of vitamin contents between buckwheat flour and some cereal flour

Vitamina Buckwheat Wheat Rice Maize Oat Sorghum

Vitamin B1 0.38 0.19 0.21 0.29 0.27

Vitamin B2 0.22 0.06 0.06 0.09 0.17 0.09
Vitamin PP 4.1 1.6 2.2 1.6 0.8 2.8
Vitamin E 1.4  0.79b
Vitamin E 0.92  0.45c
a
mg per 100 g dwb (dry weight basis).
b
Mean of 565 common buckwheat genotypes.
c
Mean of 491 Tartary buckwheat genotypes.
Source: Data from Wang LM, Zhong SH, and Cai SL (1991) Contents of vitamin PP and vitamin E in Chinese (common and tartary) buckwheat. Crop Genetic Resources 2: 2325; Wei
YM (1995) Quality and Processing of Buckwheat Grain (in Chinese). Xian, China: World Publishing Corporation.
310 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview
Fagopyritols
Two digalactosyl D-chiro-inositols and two trigalactosyl
D-chiro-inositols, members of the fagopyritol A series and
fagopyritol B series, were isolated from buckwheat (F. esculen-
tum Moench) seeds.
Fagopyrins
Fagopyrins are naturally occurring substances in the buck-
wheat plant. Its chemical structure contains a naphthodian-
throne skeleton similar that of to hypericin. Fagopyrins occur
in buckwheat grain in low quantity and are difficult to
separate.
Other Phenolic Compounds
Many other phenolic compounds (e.g., free and bound phe-
nolic acids, phenylpropanoid glycosides, anthocyanins, and
procyanidin) have been identified in buckwheat, particularly
in Tartary buckwheat. Some phenolic compounds have
been recently isolated and identified for the first time in
Tartary buckwheat, for example, 1, 3, 6, 60 -tetra-feruloyl
sucrose, 3, 6-di-p-coumaroyl-1, 60 -di-feruloyl sucrose, 1, 6,
60 -tri-feruloyl-3-p-coumaroyl sucrose, N-trans-feruloyl
tyramine, quercetin-3-O-[b-D-xyloxyl-(1!2)-a-L-rhamnoside],
and proanthocyanidins (procyanidin A).
Antinutritional Inhibitors
Antinutritional factors have been found in buckwheat grain, such
as trypsin inhibitors (I, II, and III) and tannin. They may influence
digestibility of buckwheat protein. Also, a high level of fiber in
buckwheat may be considered as an antinutritional factor.
Nutritional Value and Properties of Major Components
Protein
Buckwheat is one of the best sources of high biological value
(BV) protein in the plant kingdom. The BV of buckwheat
protein is 93, compared to egg albumin (100), soybean protein
(68), and wheat protein (63). The high BV of buckwheat
protein results in much higher utilizable protein value
(2030%) compared to other cereals, although its digestibility
is relatively lower. Buckwheat protein complements other food
proteins to improve the dietary amino acid balance. In buck-
wheat protein, threonine and methionine are the first and the
second limiting amino acids, respectively, but they are quite
abundant in other plant proteins. Compared with cereal
Table 5 Functional properties of common buckwheat protein concentrates, compared to other protein isolates
Protein concentrate Emulsifying activity at pH 7.0 (%)
Buckwheat 63.8
Soybean 50.6
Amarantha 73.3
a
From A. cruentus genotype (K350).
Source: Data from Bejosano FP and Corke H (1998) Amaranthus and buckwheat protein concentrate effects on an emulsion-type meat product. Meat Science 50: 343353.
proteins, buckwheat protein is particularly rich in lysine and
contains more arginine and aspartic acid. Buckwheat protein
also shows different bioactive effects, for example, cholesterol-
lowering effect, antihypertensive effect, antitumor activity, and
improvement in the constipation and obesity conditions by
acting in a similar way as dietary fiber and interrupting in vivo
metabolisms.
In buckwheat grain, protein albumin is the major fraction
(2842%), and glutelin (1121%) and globulin (1420%)
are next, whereas prolamin is the minor fraction (1.72.3%).
Comparatively, wheat grain is high in prolamin and glutelin
(7075%). These differences between buckwheat and wheat
lead to differences in food functional properties and processing
quality. Because of the very low level of prolamin in
buckwheat (i.e., free from gluten) and based on chemical and
immunologic studies, buckwheat protein is a valuable source
of dietary protein for gluten-sensitive individuals and can be
used for patients with gluten sensitivity (celiac disease).
Studies at the University of Hong Kong found that buck-
wheat protein exhibited much higher solubility, better emulsi-
fying activity, and slightly lower foaming ability, compared to
soy protein, and poorer foaming ability than amaranth protein
(Table 5). Japanese studies indicated that the physicochemical
properties of buckwheat protein were obviously different from
those of soy protein and casein. Buckwheat protein affected the
rheological properties of maize starch. An increase in the peak
viscosity due to the addition of buckwheat protein concentrate
was observed, but a lesser increase from the addition of buck-
wheat protein hydrolysates. From the established relationship
between pasting viscosity and granule swelling during gelatini-
zation, it was concluded that the observations were due to the
protein exerting a stabilizing factor on starch granule integrity.
It was also observed that buckwheat protein addition weak-
ened starch gel structure. This phenomenon was most likely
due to the preferential interactions between starch granules
and proteins during gelatinization and retrogradation.
Furthermore, buckwheat protein concentrate was examined
for direct utilization in an emulsion-type meat product com-
prising lean beef, pork fat, salt, and water. It was found that the
use of buckwheat protein concentrate considerably affected
both the emulsion and the cooked meat gel properties. Buck-
wheat protein showed favorable effects to be suitable as a meat
extender, as good as soy protein. Also, buckwheat protein
concentrate was tested for direct application in other food
systems (wheat dough and noodles). The addition of the buck-
wheat protein concentrate could improve the mixing proper-
ties of wheat flour and produce a positive influence on noodle
formulation and final product quality.
However, some reports pointed out a problem of the low
digestibility of buckwheat protein, which might result from
Solubility (%) Foam stability (ml)
64.2 14
33.3 20
56.5 56
 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview
antinutritional factors: trypsin inhibitors, tannin, and a high
content of dietary fiber. Buckwheat trypsin inhibitors are rela-
tively thermostable, acid-resistant, and partially susceptible to
pepsin action. In addition, allergens in buckwheat proteins
have attracted attention. During buckwheat grain milling and
food processing, some workers have allergic reactions, such as
nose itching, sneezing, rhinorrhea, dyspnea, asthma, and even
more serious symptoms. It is necessary to know more about
the behavior and processing properties of specific buckwheat
allergenic proteins.
Starch
Buckwheat starch normally has higher peak viscosity, higher
gelatinization temperature, greater swelling power, and higher
susceptibility to acid and enzymatic degradation when com-
pared with cereal starches (maize and wheat). Buckwheat
starch retrogradation is generally slower than that of maize
and wheat starch. Buckwheat starch has lower syneresis when
stored at 4  C for 310 days and has better stability to syneresis
after freezethaw cycles at 12 to 25  C.
The starch properties of common and Tartary buckwheat
compared to wheat starch. Clear differences were found in
physicochemical properties between buckwheat and wheat,
but less variation was observed between three common and
three Tartary buckwheat genotypes. Buckwheat starch had
higher peak gelatinization temperature (average Tp 68.7  C)
than standard wheat starch (Tp 63.7  C). The important past-
ing properties of buckwheat starch were higher hot paste vis-
cosity and cool paste viscosity, better resistance to shear
thinning, and less effect of NaCl on peak viscosity than for
wheat starch. Starch swelling volume is 27.428.0 ml for com-
mon buckwheat and 26.530.8 ml for Tartary buckwheat,
compared to the reference (20.1 ml) for wheat starch.
Investigations on the nutritional properties of buckwheat
starch showed that it has potential use in the design of foods
with lower glycemic index properties. The in vitro rate of starch
hydrolysis and resistant-starch formation in boiled buckwheat
groats and in a series of buckwheat/wheat breads and noodles
(3070% buckwheat flour) were evaluated. The highest level of
resistant starch (6% total starch basis) and the lowest starch
hydrolysis index were found in boiled buckwheat groats. The
rate of in vitro amylolysis was much lower in all buckwheat
products compared to the reference whole wheat bread or
noodles. Consumption of boiled buckwheat groats or bread
Table 6 Clinical effects of tartary buckwheat flour on hyperlipidemia and diabetes (hyperglycemia) after 1-month dietary treatment
Testing indexa
Hypolipidemic effect
Serum triglyceride (mg dl1) (n 18)
Serum cholesterol (mg dl1) (n 13)
Hypoglycemic effect
Blood sugar (mg dl1) (n 23)
Urinary sugar (g 24 h1) (n 27)
a
n is number of subjects.
Source: Data from Lin RF, Zhou MD, Tao YR, Li JY, and Zhang ZW (eds.) (1992) Proceedings of the 5th International Symposium on Buckwheat, 2026 August, Taiyuan, China.
Beijing: Agricultural Publishing House; Corke H and Lin RF (eds.) (1998) Proceedings of the 1st International Conference on Asian Food Product Development Focus on Specialty
Grains and Grain Products, 610 September, Taiyuan, China. Beijing and New York: Science Press.
311
and noodles containing more buckwheat induced significantly
lower postprandial blood glucose and insulin responses than
with whole wheat bread.
Additionally, there are effects of processing treatments
(including steaming or autoclaving and extruding) on buck-
wheat starch properties. The results showed that hydrothermal
treatment increases gelatinization temperature, swelling
power, and solubility of buckwheat starch. Autoclaving/cool-
ing treatment of buckwheat groats resulted in improved avail-
ability of buckwheat starch and led to more apparent amylose
and true amylose in the buckwheat starch and more resistant-
starch formation in comparison to the untreated samples. It
was also reported that extrusion influenced the physico-
chemical properties of starch and formation of starchprotein
and starchlipid complexes in extruded mixtures containing
buckwheat flour and starch. Starchprotein complexes formed
during extrusion played an important role in the stability of the
porous structure in the extruded products containing buck-
wheat starch.
Therapeutic Effects of Buckwheat
Buckwheat, especially Tartary buckwheat, has been regarded as
an effective medicinal herb in Chinese medicine for more than
1000 years. Over 80 prescriptions utilizing buckwheat in the
treatment of many diseases have been described in Chinese
traditional medicine books. In modern times, buckwheat
products are popular as functional or therapeutic foods in
China and some other countries. Regular consumption of
buckwheat foods may reduce the occurrence of hyperlipid-
emia, obesity, and diabetes. A clinical trial in Beijing Tongren
Hospital showed that Tartary buckwheat flour had hypolipi-
demic and hypoglycemic effects in treating hyperlipidemia and
diabetes (Table 6). Table 7 shows that health foods made from
Tartary buckwheat have positive effects in reducing cholesterol,
blood lipid, blood sugar, urinary sugar, and other indices.
These therapeutic effects should be attributed to unique pro-
teins, resistant starch, and dietary fiber as well as vitamins,
minerals, flavonoids, and phytosterols in buckwheat.
Protein and indigestible carbohydrates (fiber and resistant
starch) of buckwheat (particularly Tartary buckwheat) can be
used as potential functional food additives to treat conditions
such as hypertension, obesity, and constipation. Buckwheat
protein extract could lower blood cholesterol level more
Before treatment After treatment
310  241 233  173
247  57 197  43
202  46 157  40
44  43 20  22
312 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview
Table 7 Therapeutic effects of health foods made from Tartary buckwheata
Treatment Triglyceride (mg dl1) Cholesterol (mg dl1)
Buckwheat 207.8 148.1
Control 286.9 202.6
a
Statistical data from 187 samples of clinical observations by Beijing Grain Science Research Institute, Beijing. Lin RF, Zhou MD, Tao YR, Li JY, and Zhang ZW (eds.) (1992)
Proceedings of the 5th International Symposium on Buckwheat, 2026 August, Taiyuan, China. Beijing: Agricultural Publishing House; Corke H and Lin RF (eds.) (1998) Proceedings
of the 1st International Conference on Asian Food Product Development Focus on Specialty Grains and Grain Products, 610 September, Taiyuan, China. Beijing and New York:
Science Press.
efficiently, compared with soy protein and casein. Sulfur-
containing amino acids and glycine and arginine and lysine
are also considered to have important influence on this
cholesterol-lowering effect. These amino acids can conduct
downregulation of hepatic LDL receptors and improve the
removal of cholesterol and its esters in outlying tissues and
thus reduce the serum cholesterol level. Many studies have
further indicated that consumption of buckwheat protein can
cause suppression in body fat, constipation, mammary carci-
nogenesis, and colon carcinogenesis in rats and in the forma-
tion of cholesterol gallstones in hamsters. Some researchers
explain that the cholesterol-lowering, obesity-treating, and
diabetes-controlling effects are also possibly associated with
other components (dietary fiber and resistant starch) in buck-
wheat. The metabolites, formed during fermentation of resis-
tant starch and other indigestible carbohydrates in the large
intestine, contribute to the maintenance of colon health and
also have beneficial effects on glucose metabolism. Moreover, a
novel protein isolated from Tartary buckwheat water-soluble
extracts exhibited antiproliferative effect and antitumor activity.
The flavonoids from buckwheat (especially Tartary buck-
wheat) have been found to be effective in reducing blood
cholesterol levels, keeping capillaries and arteries strong and
flexible, improving blood microcirculation, and protecting
blood vessels from rupturing and forming clots. These flavo-
noids also demonstrate antioxidant, antimicrobial, and anti-
inflammatory activities. Because Tartary buckwheat leaves and
stems contain high concentrations of rutin and other flavo-
noids at flowering, extracts of buckwheat plants are helpful for
treating keratitis, while dried powdered buckwheat plants are
sometimes used for curing external wounds and ulcers.
The vitamins and minerals (including Mg, Fe, and Se) in
buckwheat can also play a crucial complementary role in the
previously mentioned therapeutic effects. Phytosterols found
in buckwheat grain, although at a low level, also showed
positive effects on blood cholesterol level. Other functions of
buckwheat sterols are waiting for further investigation, because
some phytosterols from other plant sources were found to have
strong pharmaceutical effects, for example, antivirus and anti-
tumor activities. Also, there is a report that the fagopyrins
isolated in buckwheat grain are an effective component for
the treatment of type 2 diabetes. However, there is still some
dispute on this issue.
Buckwheat Food Products
Buckwheat products are recognized as functional food in some
Asian countries. Buckwheat, as a nutritional staple crop and
Lipoprotein (mg dl1) Blood sugar (mg dl1) Urinary sugar (nmol 24 h1)
503.2 178.5 284
616.0 255.6 363
medicinal herb for Chinese consumption, has been widely
grown for a long time. A rich array of traditional buckwheat
foods are found in various places of China, such as buckwheat
wantuo and helao (or hele), buckwheat cats-ear noodles,
buckwheat noodles (vermicelli), pancake, griddle cakes, jelly
noodles, and porridge. These foods are famous and popular in
China, particularly for their long history, rich nutritional pro-
file, special flavor, various processing methods, and functional
food role in maintaining human health. For example, buck-
wheat helao (hele), wantuo, and cats-ear noodles are well-
known local foods in the northern region of China (Shanx
and Shaanxi provinces).
In Japan, buckwheat noodles called soba and buckwheat
roasted groats have long been very popular, while in Korea, a
favorite jelly-type food called mook is also made from buck-
wheat. In India, buckwheat flour is used for making chillare,
an unleavened bread fried with ghee, and also for a crisp food
called pakora. Honey produced from common buckwheat is
prized in both China and India. In eastern Europe, buckwheat
grain forms an important part of the traditional diet. Buck-
wheat groat porridge and meals and soup made from flour are
still favorite traditional foods in Poland, Russia, and some
other eastern European countries. In the United States,
although buckwheat is mainly used for animal feed, buck-
wheat cake was once a special breakfast item and buckwheat
flour is also an ingredient of some pancake mixes.
Modern buckwheat food products include buckwheat dried
noodles or vermicelli, instant noodles, spaghetti or macaroni,
biscuits, breads, cakes, pastry, breakfast cereals and snacks,
vinegar, drinks, and beverages (Tartary buckwheat tea, buck-
wheat beer, and liquor). Most of these products, particularly
made from Tartary buckwheat, are popular healthy foods in
China. It is estimated that commercial food products made
from buckwheat have produced over 10 billion RMB of value
in China. In addition, tender buckwheat shoots are used as
fresh vegetables and are also processed into canned vegetables.
Buckwheat Processing and Development
Buckwheat grains are dehulled and milled into flour for food
processing. Buckwheat flour milling is similar to wheat dry
milling. Buckwheat flour yield ranges from 58% to 75%,
mainly depending on variety and milling techniques. Buck-
wheat flour contains similar levels of starch and protein to
wheat flour. However, the behavior of buckwheat flour is
different from wheat flour, and it is difficult to make wheat
flour-like foods because there is no gluten. Therefore,
 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview 313

buckwheat flour is usually mixed with wheat flour and other
flours before use.
Chinese researchers and food factories have mixed buck-
wheat and wheat flour along with other flours to successfully
make buckwheat pastry, sandwich cake, dried noodles, instant
noodles, and spaghetti (Tables 8 and 9). The processing steps
for dried noodles are shown in Figure 1. Suitable levels of
buckwheat flour are
10% for crisp pastry and cakes,
2040% for noodles, and 3050% for palatable spaghetti.
The cooking quality of spaghettis mixed from different ratios
of buckwheat and wheat flours is shown in Table 9. Buck-
wheat/wheat spaghettis generally have higher water absorp-
tion, higher cooking loss, and higher protein loss rate than
pure wheat spaghetti, because buckwheat flour contains much
more albumins (water-soluble) and globulins (soluble in salt
water) than wheat flour. These commercial foods are produced
by modern food-processing equipment. However, most tradi-
tional buckwheat foods are still handmade in the countryside
and even cities in China. An example of the basic hand-
preparation method for buckwheat helao is shown in Figure 2.
In Japan, there are many noodle-making factories produc-
ing soba noodles. Buckwheat flour used for making soba is
commonly mixed with 1050% wheat flour. These noodles
are sold in precooked, boiled, dried, or instant stages, as well
as being made for direct sale to buckwheat noodle restau-
rants, small shops, and stands. In Europe and the United
States, buckwheat flour is mixed with other cereal flours to
make pancakes, flakes, biscuits, bread, noodles, spaghetti or
macaroni, and ready-to-eat breakfast cereals. The processing
of these foods can be conducted using ordinary baking,
extruding, or expansion methods. The amount of buckwheat
flour added to other cereal flours is generally < 50%. For
instance, acceptable bread contained 25% buckwheat flour
with a formulation that included vital gluten, whey, or sour
milk. A recipe for bread with 10% Tartary buckwheat flour is
shown in Table 8. It was also reported that the buckwheat
corn flakes were produced by extrusion processing. The for-
mulations with an addition of 18%, 22%, and 33% buck-
wheat flour were used. The addition of 30% buckwheat flour
gave the best organoleptic properties.
Although buckwheat is still cultivated as a minor crop in the
world, buckwheat product development has quickened since
the 1990s, particularly in China. To date, there has been an
emerging trend in the research and development of high value-
added buckwheat products, especially Tartary buckwheat
products. Tartary buckwheat has high potential in the health
food market. Since Tartary buckwheat contains a high level of
flavonoids and other therapeutic components, some cosmetics
and medicine products have successfully been developed and
marketed using buckwheat materials in China, such as cream,
shampoo, lotion, toothpaste, and flavonoid capsules. How-
ever, many such new products have not been accepted well in
the marketplace, because of their high cost and erratic quality
control.

Table 8 Recipes for three selected buckwheat baked foodsa

Materials (g) Chinese crisp pastry Sandwich cake Mixing bread

Buckwheat flour 1.0 1.0 25b

Wheat flour 8.5 8.5 225
Soybean flour 0.5 0.5
Egg 10
Shortening oil 3 10
Salt 0.01 3.75
Sugar 8.0 10
Refined maltose 1.0 1.6
Skimmed milk 5
Dried yeast 3
Flavor (prickly ash) 0.10.15
Jam 3
Buckwheat flour paste 0.5
a
In these selected foods, buckwheat flour accounts for 10% of total composite flour in the baking foods: data from Lin RF, Zhou MD, Tao YR, Li JY, and Zhang ZW (eds.) (1992)
Proceedings of the 5th International Symposium on Buckwheat, 2026 August, Taiyuan, China. Beijing: Agricultural Publishing House.
b
Tartary buckwheat flour.

Table 9 Cooking quality of spaghetti made from common buckwheat and wheat flour at different ratios

Loss of protein (%)

Buckwheat/wheat flour ratio Water absorption (%) Cooking loss of dry materials (%) Cooking in water Cooking in salt water

0:100 218 6.4 1.4 6.6

25:75 235 6.6 2.0 7.5
50:50 251 8.2 3.7 13.9
75:25 278 8.5 2.7 9.4

Source: Data from Wei YM (1995) Quality and Processing of Buckwheat Grain (in Chinese). Xian, China: World Publishing Corporation.
314 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview

Buckwheat and wheat flour (3 : 7) Buckwheat grains

Mix flour De-hull and stone-mill into flour

Add ~40 C water (1% NaCl) (H2O : flour 3.5 : 6.5)

Add water (water : flour 4 : 6) and a little limewash

Knead dough for 20 min and

ripe for 20 min Mix and knead well into soft and
smooth dough (light green)

Roll (1.21.4 mm thickness) and slice (1.51.8 mm width)

Press dough into rough noodle shape by helao instrument

Dry at <40 C for 78 h

Fresh noodle strands are directly

dropped into boiling water
Cut (20 cm length)

Cook at ~ 100 C for 1015 min

Final product of dried noodles

Figure 1 Processing procedure for dried noodles of common or Tartary

buckwheat.
Buckwheat helao

Nowadays, buckwheat functional foods and related prod- Figure 2 Traditional method of buckwheat helao preparation.
ucts have attracted widespread interest and have become more
popular. Much more attention should be paid to the biological
functions, palatability, and acceptability of buckwheat prod-
into added-value components that would be attractive to
ucts. It is necessary to enhance development and commercial-
the health food industry.
ization of traditional buckwheat foods and to improve modern
buckwheat food-processing techniques.
Compare Chinese and Japanese buckwheat noodles in
depth, showing how different traditional products achieve
their required quality outcomes. Suggest new product ideas
for buckwheat-based noodles.
Exercises for Revision

Describe the composition of buckwheat seed in relation to
physical properties of the food product (e.g., the effect of
starch on noodle texture).
Outline the health claims made for the various fractions
and components of the buckwheat seed.
Try to make buckwheat noodles, experimenting with the
effect of varying the proportion of wheat flour to buck-
wheat flour.
See also: Carbohydrates: Starch: Chemistry; Food Grains:
Intolerance, Allergy and Diseases: Celiac Disease; Proteins:
The Protein Chemistry of Cereal Grains; The Basics: Taxonomic
Classification of Grain Species; The Legumes and Pseudocereals:
Amaranth: Overview; Pseudocereals: Overview.

Further Reading
Bejosano FP and Corke H (1998) Amaranthus and buckwheat protein concentrate effects
Exercises for Readers to Explore the Topic Further on an emulsion-type meat product. Meat Science 50: 343353.
Corke H and Lin RF (eds.) (1998) Proceedings of the 1st International Conference on
Design a breeding program for the development of nutri- Asian Food Product Development Focus on Specialty Grains and Grain Products,
tionally improved buckwheat, suitable for fractionation Taiyuan, China, 610 September. Beijing and New York: Science Press.
 THE LEGUMES AND PSEUDOCEREALS | Buckwheat: Overview
Guo XN, Zhu KX, Zhang H, and Yao HY (2010) Anti-tumor activity of a novel protein
obtained from tartary buckwheat. International Journal of Molecular Sciences
11: 52025212.
Joshi BD and Rana RS (1995) Buckwheat (Fagopyrum esculentum). In: Williams JT
(ed.) Cereals and Pseudocereals, pp. 85127. London: Chapman and Hall.
Li WD, Lin RF, and Corke H (1997) Physicochemical properties of common and tartary
buckwheat starch. Cereal Chemistry 74: 7982.
Li SQ and Zhang QH (2001) Advances in the development of functional foods from
buckwheat. Critical Reviews in Food Science and Nutrition 41: 451464.
Lin RF (1994) Buckwheat in China. (in Chinese). Beijing: Chinese Agriculture
Publishing House.
Lin R, Li W, and Corke H (1998) Spotlight on Shanxi province China: Its minor crops
and specialty foods. Cereal Foods World 43: 189192.
Lin RF, Zhou MD, Tao YR, Li JY, and Zhang ZW (eds.) (1992) Proceedings of the 5th
International Symposium on Buckwheat. Taiyuan, China, 2026 August. Beijing:
Chinese Agricultural Publishing House.
Ren Q, Wu CS, Ren Y, and Zhang JL (2013) Characterization and identification
of the chemical constituents from tartary buckwheat (Fagopyrum tataricum
Gaertn) by high performance liquid chromatography/photodiode array
detector/linear ion trap FTICR hybrid mass spectrometry. Food Chemistry
136: 13771389.
315
Tao CJ, Yuan P, Wang X, and Liao B (2014) Advances in the development of buckwheat
products. Cereal and Food Industry 21: 5254.
Wang LM, Zhong SH, and Cai SL (1991) Contents of vitamin PP and vitamin E in
Chinese (common and tartary) buckwheat. Crop Genetic Resources 2: 2325.
Wei YM (1995) Quality and Processing of Buckwheat Grain (in Chinese). Xian, China:
World Publishing Corporation.
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Chinese). Beijing: Chinese Science Press.
Relevant Websites
http://www.buckwheatgrowers.com Information on buckwheat growers, products and
services.
https://www.hort.purdue.edu/newcrop/afcm/buckwheat.html Center for New Crops
and Plant Products at Purdue University. New CROP provides windows to new and
specialty crop profiles, including buckwheat.
 Chickpea: Overview
EJ Knights and KB Hobson, New South Wales Department of Primary Industries, Tamworth, NSW, Australia
2016 Elsevier Ltd. All rights reserved.
This article is a revision of the previous edition article by E.J. Knights, volume 1, pp 280287, 2004, Elsevier Ltd.
Topic Highlights
Chickpea is an annual legume belonging to the Cicer genus.
Seed shape separates the species into three main groups:
desi, kabuli, and pea.
Chickpea is a key dietary component in many countries
where vegetarian diets dominate or animal protein is too
expensive.
Chickpea is grown mainly as a sole crop in a range of
rotations.
World production averages 10 Mha annually.
India produces more than 67% of the worlds annual
production.
Learning Objective
To achieve understanding of the agriculturally important
species in both farming systems and its end uses
Introduction
Chickpea (Cicer arietinum L.) is a herbaceous, annual legume.
Plants are typically short (2050 cm) with a semierect to erect
growth habit and a variable number of primary and secondary
branches. An imparipinnate leaf type, producing 1118 small
leaflets, gives the foliage its distinct fernlike appearance
(Figure 1). Chickpea is a long day plant; however, time to
flowering is accelerated by increasing temperature. Flowering
is indeterminate with single (rarely double), small purple or
white flowers produced on racemes arising from the leaf axil.
Pods have a characteristic oval to rhomboid shape and inflate
rapidly after (self-) fertilization (Figure 1). All aerial plant parts
excepting flowers have a dense covering of fine hairs, which
secrete a mixture of organic acids. Roots are colonized by
Mesorhizobium ciceri, a nitrogen-fixing bacterium specific to
the genus Cicer. Nodules formed by these bacteria vary in
size, the largest approaching 3 cm in diameter. The seed is
highly variable in appearance but invariably possesses the
characteristic chickpea beak protruding over the embryo.
Seed shape, color, and, to a lesser extent, size separate the
species into three main groups: desi, kabuli, and pea
(Figure 2).
Desi (derived from the Hindi and Urdu word for native or
local) seeds have an angular shape and thick, colored (mostly
brown) seed coat. In contrast, kabuli seeds have a more
rounded shape and thin, white to cream-beige seed coat.
There is considerable overlap between these two seed types in
seed size; however, desi seeds are generally smaller (weighing
80350 mg) than kabuli seeds (100750 mg). The pea-type
seed has little commercial significance.
316 Encyclopedia of Food Grains, Second Edition
Origin and Spread
The genus Cicer contains 43 species, 34 perennial and 9
annual. All annual species are diploids and have 16 chromo-
somes (2N 16). Among these, C. reticulatum is the most
closely related to the cultivated chickpea and the presumed
progenitor. Domestication occurred about 10 000 years ago
in the region of southeastern Turkey, part of the Fertile Cres-
cent. Recent evidence suggests this occurred as a single event
due to the limited geographic distribution of the wild species
populations and the very low genetic variation of the cultigen.
About 40005000 years ago, the domesticated forms, initially
desi types, commenced a westward movement along estab-
lished trade routes toward the Mediterranean. Subsequent
spread to southern Europe and North Africa, including Ethio-
pia, occurred by land and/or sea routes. Differentiation of the
kabuli type is thought to have occurred in the Mediterranean
region in comparatively recent times. The eastward movement
of desi types to India commenced about 4000 years ago,
whereas kabuli types did not arrive there until about 300
years ago. (The prominence of Kabul on the Silk Road sug-
gests both an overland introduction and the origin of the word
kabuli.) In the sixteenth century, Spanish and Portuguese
travelers carried kabuli types to South and Central America
where they came to be known by their Spanish name gar-
banzo. Chickpea industries have comparatively recent histo-
ries in the United States (1950s), Australia (1970s), and
Canada (1990s).
Production
During the 1990s, world chickpea production has averaged
8 million tonnes (Mt) from 10.5 million hectares (Mha). In
the 2000s, this increased to 8.5 Mt from a similar area, with the
increased production coming from increased yield. Crop area
for the past 3 years (20112013) has averaged 13 Mha, which
is the highest 3-year average production (Figure 3). Histori-
cally, Asia has been the major chickpea producer. For years
20092013, it accounted for 85.1% of all production, followed
by Australia (5.3%), Africa (5.0%), Americas (3.8%), and
Europe (0.9%). Desi types contribute 70% of production
worldwide. Chickpea is cropped in approximately 80 coun-
tries. India produces two-thirds of the world crop (Table 1),
and after a steady decline in the 1990s, production has been
increasing in recent years. A similar trend has occurred in
Pakistan, the worlds second largest producer. In European
countries, chickpea production has fallen dramatically, by
6097% since the 1960s. On the other hand, production has
risen significantly in some countries. In Australia, the worlds
fourth largest producer, chickpea area increased rapidly during
the 2000s as the demand increased for cereal break crops and
alternatives were sought for nitrogen fertilizer. Expanded chickpea
http://dx.doi.org/10.1016/B978-0-12-394437-5.00035-8
 THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview
Figure 1 Chickpea branch showing typical leaf and pod shape.
(Chickpea normally produces single, axillary flowers; the double-podded
variant shown here may confer some yield advantage in low-productivity
environments.)
Figure 2 Seed type in chickpea showing, clockwise from top of picture:
kabuli, desi, and pea (intermediate) forms.
areas in African countries such as Ethiopia, Malawi, and Tanzania
have also helped stabilize or even increase world production in
recent years. Average world yield over the period 20092013 was
925 kg ha1. Central and North America had the highest yields
(1772 kg ha1) followed by Europe (1529 kg ha1), Australia
(1222 kg ha1), Africa (1076 kg ha1), and Asia (883 kg ha1).
Yields were generally highest in countries where all or most of the
crop was irrigated, such as Israel (4723 kg ha1), Lebanon
(2592 kg ha1), and Egypt (2071 kg ha1).
Consumption and Trade
Chickpea is an important part of the diet in most producing
countries and is gaining in popularity elsewhere. Per capita
consumption is generally much higher in producing countries,
317
such as India (5.1 kg pc) and Pakistan (4.4 kg pc), than in
developed, nonproducing countries such as the United Kingdom
(0.2 kg pc). Consumption is also comparatively high in some
wealthier, producing countries such as Turkey (6.3 kg pc) and
Spain (2.3 kg pc). Historically, most chickpeas were consumed in
their country of origin. Until the late 1970s, international trade
accounted for only 2% of world production. This percentage rose
to 9% in the following three decades as demand exceeded supply
in major producing countries, especially in east Asia. During
the period 20082011, India was the largest importer
(183 000 tonnes) followed by Pakistan (177 000 tonnes) and
Bangladesh (129 000 tonnes). For the same period, Australia
was the largest exporter (414 000 tonnes), followed by India
(154 000 tonnes) and Mexico (104 000 tonnes). Indias position
as the second largest exporter of chickpea is driven by a rapid
increase in the production of large kabuli types. Globally, the
volumes of desi and kabuli types now traded are similar. Prices
fluctuate according to supply and demand; however, there is
generally a premium for kabuli types and particularly for those
with large seed (diameter 911 mm).
Role in Farming Systems
Chickpea is grown in a range of environments encompassing
extensive variation in latitude (53 N to 39 S), altitude, soil
type, photoperiod, temperature, and rainfall. Seven broadly
different regions can be distinguished.
Indian Subcontinent (India, Pakistan, Nepal,
Bangladesh, and Myanmar)
Desi types predominate in this region; however, the produc-
tion of kabuli has expanded rapidly in recent years. Chickpea is
typically grown as a winter crop with sowing from October to
December and harvest from February to April. The crop is
mostly rainfed, relying essentially on soil moisture conserved
from monsoon rains; there is little effective growing season
rainfall. Soil types vary from the heavy, black cotton soils of
central and southern India to the alluvial soils of the Indo-
Gangetic Plain and the much lighter textured sands of Rajas-
than in India and the Thal area of Pakistan. There has been a
large shift in chickpea area from north to south over the past
four decades.
Chickpea is grown mainly as a sole crop in a range of
rotations. Where sufficient rainfall or irrigation allows, for
example, in northwestern India or in the rice-based systems
of northeastern India, Bangladesh, and Myanmar, chickpea
follows rainy season crops such as rice, millet, sorghum,
maize, sugarcane, cotton, guar, and sesame. However, in
lower rainfall zones where only one crop per year is possible,
chickpea is grown as a fallow crop. Less frequently, it is inter-
cropped in varying proportions with Brassica spp., linseed,
safflower, or sorghum or sown as a mixed crop with wheat,
barley, linseed, or sorghum.
Fertilizer and pesticide use is generally low and there is
minimal mechanization of sowing, harvest, or weed, insect,
and disease control. Traditionally, these operations are per-
formed by hand or with the assistance of animal drawn
318 THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview

16000 1000
area
production
yield 900

800
Area (x1000 ha)/production (x1000 t)
12000
700

600

Yield (kg ha-1)

8000 500

400

300
4000
200

100

0 0
1962

1965

1968

1971

1974

1977

1980

1983

1986

1989

1992

1995

1998

2001

2004

2007

2010

2013
Figure 3 World area, production and yield of chickpea, 19612013. Each datum is the mean for a 3-year period. Source: FAO.

Table 1 Mean area, production, and yield of chickpea in major chickpea-producing countries (20092013)

Country Area (ha) Production (tonne)a Yield (kg ha1)

India 8 634 000 7 858 400 (67.8) 910

Pakistan 1 049 660 568 000 (4.9) 543
Iran (Islamic Republic of) 549 175 275 385 (2.4) 502
Australia 509 162 609 402 (5.3) 1222
Turkey 437 471 520 935 (4.5) 1191
Myanmar 326 106 461 708 (4.0) 1414
Ethiopia 202 927 333 377 (2.9) 1673
Malawi 115 545 61 075 (0.5) 526
Mexico 92 938 163 674 (1.4) 1705
United Republic of Tanzania 77 083 74 425 (0.6) 938
Syrian Arab Republic 76 944 52 749 (0.5) 685
a
Percentage of world production in brackets.
Data from FAO.

implements, although some regions are introducing mechani- mainly on growing season rainfall. Winter sowing occurs in
zed processes. Egypt and Sudan. Sowing is usually delayed until late February
to May to minimize losses from cold and/or ascochyta blight.
Rapidly increasing temperatures and moisture stress in summer
West Asia, North Africa, and Southern Europe hasten maturity and limit crop duration to 90120 days.
Chickpea is generally confined to areas having an annual
Production in this region is almost exclusively of kabuli types. rainfall above 400 mm. It is grown as a sole crop in rotation
The crop is grown at high altitudes in west Asia (Afghanistan, with wheat (bread or durum) or barley; a fallow phase or
Iran, and Turkey) but at comparatively lower altitudes in the summer or forage crop may be included in drier areas. There
Middle East (Syria, Yemen, Jordan, Israel, and Iraq), North is some mechanization of production, particularly sowing and
Africa (Egypt, Sudan, Algeria, Tunisia, and Morocco), and south- weeding, but harvest is still largely manual. High labor costs
ern Europe (Greece, Italy, Spain, and Portugal). It is grown as a have contributed to the decline in chickpea production in parts
spring crop in this Mediterranean-type environment, relying of the region, especially in Europe.
 THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview
East Africa (Ethiopia, Eritrea, Kenya, Tanzania, Malawi,
and Uganda)
Mostly desi and some kabuli chickpeas are grown in this
region, usually at high altitudes (up to 2400 m). The area of
kabuli is increasing in some countries with the introduction of
improved varieties and is up to one-third of the crop in Ethi-
opia. Crops are mainly sown as pure stands in rotation with
wheat, barley, and teff, but to a lesser extent as mixtures with
sorghum, safflower, and maize. Crop growth relies mainly on
residual soil moisture. Sowing follows the wet season, from
July to early September in Ethiopia and from February to April
further south. Inputs of fertilizers and pesticides are minimal,
and although there is some use of tractors in sowing, harvest
and threshing remain manual operations.
Central and Southern America (Mexico and Argentina)
In Mexico, desi chickpeas are produced under rainfed condi-
tions in the west-central parts of the country and kabuli types
under irrigation in the drier northwest. Chickpea is grown as a
winter crop following maize in the west-central region and
following soybean or sesame in the northwest. Sowing is from
October to December and harvest from March to May. There
has been some recent transition to mechanized production,
particularly for kabuli types. Production in Argentina has
recently increased to 40 000 ha of kabuli type. It is sown
from April to July.
North America (Canada and the United States)
The North American chickpea industry began in coastal dis-
tricts of California where spring-sown kabuli crops were grown
on residual soil moisture. Later, winter crops were grown under
irrigation in the drier San Joaquin Valley. During the 1980s,
both desi and kabuli chickpeas were introduced to the Palouse
region of northwestern United States and, in the following
decade, to the brown and dark brown soil zones of Saskatch-
ewan and Alberta in Canada. Current production areas are the
Pacific Northwest in the United States and the province of
Saskatchewan in Canada. The threat of cold and ascochyta
blight forces chickpeas to be grown as a spring crop in both
these regions. Sowing commences in May and harvest, partic-
ularly in Canada, needs to be completed by September to avoid
freezing injury to immature seeds. In the United States and
Canada, all stages of chickpea production, including seed
inoculation; sowing; weed, disease, and insect control; harvest;
and seed drying, are highly mechanized. In most cases, the
machinery and processes have been modified from those
used in cereal production.
Australia
National production (90%) is mainly desi type and this
predominantly occurs in the summer rainfall-dominant
regions in the northeast of Australia. In the winter
rainfall-dominant Mediterranean-type environments in the
319
southeast and southwest, production in the southeast region
is mainly kabuli type. Chickpea is grown as a winter crop,
sown from May to July and harvested from October to January.
Most crops are produced under rainfed conditions; incrop
rainfall is a major yield determinant in most regions, especially
in the southeast and southwest. As in North America, produc-
tion is highly mechanized. Control of ascochyta blight, the
major industry problem in most regions, relies heavily on
ground rig application of foliar fungicides. In northeast Aus-
tralia, Phytophthora root rot is a common biotic constraint,
particularly in seasons conducive to the disease.
Production Constraints
There is a large discrepancy between the yield in farmers fields
and that commonly obtained under experimental conditions.
The low yields generally obtained by farmers reflect a plethora
of problems that beset the crop. The major production con-
straints and their impacts are described below.
Physical Constraints
Drought and Heat
Chickpea is mostly grown in low rainfall environments. Inad-
equate soil moisture is therefore a critical production con-
straint and may occur throughout the growth cycle. Winter/
spring rains in west Asia, or soil moisture remaining from the
wet season in the Indian subcontinent and East Africa, may not
be sufficient for farmers to sow; for many regions, variability in
main season rainfall explains much of the year-to-year varia-
tion in chickpea area. Receding soil moisture, particularly in
low rainfall areas in India and Pakistan, may also be inade-
quate to sustain seedlings. Intermittent moisture stress of
established crops can result from breaks in winter and early
spring rainfall, but a more serious problem is terminal mois-
ture stress suffered by spring-sown crops (and winter crops in
the Indian subcontinent) as podding coincides with rapidly
declining soil moisture. Terminal drought is a major constraint
in an estimated 80% of the global chickpea area.
Heat stress (temperature >35  C) during the reproductive
phase has become a serious constraint as chickpea moves from
cooler long-season environments to warmer short-season envi-
ronments, an increased area under later sowing, and expected
increases in temperatures due to increased climate variability.
Cold
Low temperatures affect the plant in two main ways: tissue
damage through disruption of cell membranes and abortion of
flowers. Sensitivity to low temperatures (<5  C) is one reason
for spring sowing in higher elevation areas of west Asia. How-
ever, genotypes have now been identified that can survive tem-
peratures as low as 25  C. Winter sowing of these cold-tolerant
genotypes can increase yield potential by 70% through
increased biomass production and improved water-use effi-
ciency. Where the crop is sown in autumn/winter, such as in
northern India, Pakistan, and Australia, chilling damage in the
early flowering phase narrows the window for pod setting.
Sowing is normally delayed so that commencement of
320 THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview
flowering coincides with mean daily temperatures favorable for
podding (15  C).
Hostile Soils
A range of soil chemical and physical problems limit chickpea
growth. The most important of these is salinity, since chickpea is
comparatively intolerant of the ionic imbalances characteristic of
saline soils. Sodic soils, in which high levels of exchangeable
sodium are associated with increased bulk density, low water
infiltration, and poor aeration, are also common in northern
India and in other regions where soil pH is very high. Retarded
root growth and poor nodulation inhibit chickpea growth and
productivity under such conditions. Other locally important soil
factors are acidity (poor nodulation), deficiencies of iron and
zinc, and either a deficiency or toxicity of boron.
Diseases
Ascochyta Blight
Ascochyta blight is the most important disease of chickpea
worldwide and has been recorded in nearly all producing coun-
tries. Recent epidemics in the United States (1980s) and Austra-
lia (1990s) caused major industry disruptions, reflected by a
sharp decline in the area sown. Elsewhere, for example, in west
Asia and the Mediterranean region, the threat of ascochyta blight
causes sowing to be delayed, thereby significantly reducing yield
potential. The causal agent of ascochyta blight is Ascochyta rabiei
(Passerini) Labrousse (syn. Phoma rabiei and teleomorph Didy-
mella rabiei (Kovachevski)), a fungus spread by infected seed and
crop residues, long-range dispersal of sexually produced asco-
spores, or, within the crop, short-range dispersal of asexually
produced pycnidiospores. Lesions are formed on leaves, stems,
pods (Figure 4), and seeds. The disease progresses quickly under
favorable wet conditions and can cause total crop failure. Asco-
chyta can be controlled by an integrated disease management
approach based on host resistance and, depending on location,
removal of infected plant debris, delayed sowing, seed dressing,
and foliar fungicides.
Other Foliar Diseases
Botrytis gray mold (Botrytis cinerea Pres.) is the most important
of the foliar diseases after ascochyta. The pathogen has a very
broad host range; therefore, the disease is widely distributed.
Most damage occurs under high humidity and generally at
temperatures higher than optimum for ascochyta. Botrytis
has contributed to the decline of the Bangladesh chickpea
industry and has also caused significant or total crop losses
in the higher rainfall zones of northeastern India and in Aus-
tralia. The following diseases are also regarded as locally sig-
nificant: sclerotinia stem rot (Sclerotinia sclerotiorum (Lib.)
de Bary), phoma blight (Phoma medicaginis Malbr. and
Roumeguere), alternaria blight (Alternaria alternata (Fries:
Fries) Keissler), stemphylium blight (Stemphylium sarciniforme
(Cavara) Wiltshire), and rust (Uromyces ciceris-arietini Jaczewski
in Boyer & Jaczewski).
Figure 4 Ascochyta blight on pods. The concentric rings of dark-
colored pycnidia (fruiting bodies) are a useful diagnostic feature for this
disease in chickpea.
Fusarium Wilt
Fusarium wilt is the other major disease of chickpea and, like
ascochyta blight, causes economic damage in a large number of
producing countries. It is a soil- and seed-borne disease caused
by different races of the fungus Fusarium oxysporum f.sp.ciceris
Matuo and K. Sato. The races are differentiated on their ability
to incite specific reactions causing two types of symptoms:
yellowing syndrome and wilting syndrome. The wilting syn-
drome is more destructive and economically more important
than yellowing syndrome. Fusarium wilt is more frequently
expressed under warm, dry growing conditions and is therefore
a greater problem in the Indian subcontinent and in regions
where the crop is spring-sown. Fungicidal seed dressings pro-
vide protection against seed-borne infection, but the longevity
of the pathogen in soil makes host resistance the most success-
ful way of combating the disease.
Root and Collar Rots
A number of fungal root or collar diseases have the potential to
inflict significant yield losses. Dry root rot (Rhizoctonia batati-
cola (Taubenhaus) Butler) is the most important of these,
especially on the Indian subcontinent. The disease generally
presents as a sudden drying of scattered plants at the podding
stage and is favored by hot, dry conditions. Other diseases
include wet root rot (Rhizoctonia solani Kuhn), black root rot
(Fusarium solani (Martius) Saccardo), collar rot (Sclerotium rolf-
sii Saccardo), and phytophthora root rot (Phytophthora medica-
ginis Hansen and Maxwell). The latter has been recorded in a
number of countries, but economically significant damage has
only been observed in northeastern Australia where it is the
major production problem.
Viruses
Considerable crop losses have been attributed to virus disease,
especially in India, Pakistan, Iran, the United States, and
Australia. In most cases, a complex of viruses has been
 THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview
implicated. Aphids, particularly Aphis craccivora Koch, are almost
always the vector responsible for disease transmission. In India,
chickpea stunt is the name given to a syndrome characterized by
foliage discoloration (red in desi and yellow in kabuli), stunting,
phloem browning, and plant death. The disease has been
ascribed to the leafhopper-transmitted chickpea chlorotic
dwarf virus and some aphid-transmitted luteoviruses, including
bean leafroll luteovirus. Other viruses known to cause disease
are alfalfa mosaic virus, cucumber mosaic virus, beet western
yellows virus, and subterranean clover red leaf virus.
Nematodes
Nematodes reduce yield by colonizing and damaging roots,
reducing the efficiency of nitrogen fixation and in some cases
acting synergistically with fungal diseases such as fusarium
wilt. Many nematode species are known to infect chickpea
although only three are considered economically important.
Root-knot nematodes (Meloidogyne spp.) cause abundant for-
mation of galls, whereas both cyst nematode (Heterodera ciceri
Vovlas, Greco & di Vito) and root-lesion nematodes (mainly
Pratylenchus thornei Sher and Allenand and P. neglectus
(Rensch) Chitwood and Oteifa) damage root cellular tissue
causing necrotic regions. Root-knot nematodes and root-lesion
nematodes have an extensive host range and are widely dis-
tributed; cyst nematode has a very limited host range and its
distribution is confined to the Middle East.
Insects
The small number of insect pests causing economic damage to
chickpea is attributed to the antibiotic effect of the plants
acidic (mainly malic acid) exudate. The most widespread and
damaging pest is the pod borer (Helicoverpa spp.). Various
larval instars feed on leaves, flowers, and pods. Most yield
loss is caused when late instar larvae enter the pod and partly
or fully consume the developing seed. There is little effective
host resistance within chickpea so control is reliant on a com-
bination of chemical and biopesticides (such as nuclear poly-
hedrosis virus). Genetic transformation has been used to
develop chickpea containing the toxins from the bacterium Bt
and it is currently undergoing field testing.
The other major field pest is leaf miner (Liriomyza cicerina
(Rondani)) whose distribution is confined to the Mediterra-
nean region. Larvae tunnel through the leaf parenchyma caus-
ing loss of photosynthetic tissue and premature leaf drop.
Infestation of stored seed is mainly by small bruchid beetles
(Callosobruchus spp.). Larvae emerge from eggs laid on the
(stored) seed surface and tunnel beneath the seed coat where
they feed and pupate. Adults emerge from the seed, ready for
the next life cycle and leaving a substantial cavity.
Weeds
Chickpeas slow early growth and low total biomass predispose
the crop to severe weed competition. In most countries, weed
control is done by manual weeding or cultivation by animal-
or tractor-drawn implements. These practices are feasible
where crop area is small, although yield potential is often
compromised where wide rows are employed to facilitate
321
cultivation. Options for pre- and postemergent control of
both grass and broadleaf weeds are now available and used
extensively in Australia, the United States, and Canada. Grass
herbicides are very effective, have no toxicity to chickpea, and
contribute to the productivity of ensuing cereal crops by elim-
inating graminaceous diseases that can otherwise survive on
grass weeds. The development of herbicide-tolerant varieties
will also provide further weed control options.
Utilization
Nutritional Status
Desi chickpea seeds are composed of the embryo (1.5%), seed
coat (15.5%), and cotyledons (83%); the seed coat fraction of
kabuli seeds is much lower (6.5%), resulting in a higher cotyle-
don fraction (92%). Protein averages 23% and is deficient in the
sulfur-containing amino acids methionine and cystine. Carbo-
hydrate, the main constituent, averages 63.5% of total seed and
is mainly starch. Soluble sugars comprise 10% of the starch
and include oligosaccharides such as stachyose and raffinose
that cannot be broken down by human digestive enzymes.
Gases produced by bacterial degradation of these oligosaccha-
rides cause flatulence and intestinal discomfit. Oils comprise
5% of the seed but are well below commercially extractable
levels. Chickpea is generally low in antinutritional factors such
as protease and amylase inhibitors, whose activities are further
reduced by cooking, and protein-binding phenolics, which are
largely removed with processing.
Uses and Processing
Chickpea is a key dietary component in many countries where
animal protein is too expensive or in other countries where the
crop has a long history of cultivation and consumption. The
complementarity of pulse and cereal proteins has given rise to
many traditional dishes in which chickpea is combined with
wheat, rice, or some locally important cereal, such as teff, to
provide the bulk of dietary protein and calories. The use of
chickpea as human food is hugely varied, reflecting both the
crops antiquity and its broad cultural base (Table 2). The
mature seed, either whole or processed, accounts for nearly
all consumption. However, there are specialty uses for the
green, immature plant. Fully podded plants are commonly
sold in street markets in India and the Middle East. The seeds
are either roasted in the pod, consumed raw as a snack, or
boiled; fresh leaves are also used as a vegetable. Ripe seeds (desi
and kabuli) are presoaked and boiled, with or without the
addition of vegetables or meat, to produce a range of tradi-
tional dishes. Kabuli seeds are also canned after cooking, but a
more important use is as hommos bitihneh (pureed and mixed
with oil). Both desi and kabuli seeds are also roasted. This
process generally requires a quick soaking followed by high-
temperature treatment (e.g., at 250  C) for 23 min in pre-
heated sand or an oven. Variations to the soaking, roasting,
and final processing (e.g., decortication, polishing, or sugar
coating) give this snack food its regional characteristics.
Most desi seeds are milled to produce dhal, a process that
involves removal of the seed coat and cleaving of the cotyle-
dons (Figure 5). Milling of chickpeas is a major industry in the
322 THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview

Table 2 Main uses of chickpea for human consumption

Plant/seed part used Process/product Country/region

Green leaves Boiled as vegetable Indian subcontinent (ISC)

Immature plant Roasted, pods shelled ISC, Middle East, Ethiopia
Unripened pods Pods shelled, seeds raw or boiled ISC, Ethiopia, Turkey
Whole seed kabuli Soaked, boiled alone or with vegetables, rice, meat, spices Most countries
Soaked, boiled, canned North America, Europe, Australia, Middle East
Soaked (briefly), roasted Middle East, North Africa
Soaked, boiled, pureed (hommos) Middle East, North Africa
Whole seed desi Soaked, boiled with vegetables, rice, spices ISC
Soaked (briefly), roasted ISC
Germinated ISC
Dhal Soaked, boiled with spices ISC
Flour (besan) Batter for fried vegetables ISC
Batter with rice, fermented ISC
Thickener for gravies ISC
Fried extrusion products ISC
Mixed with wheat flour, bread ISC, Middle East

countries despite the new varieties being released by national

Figure 5 Dhal produced from desi seeds following removal of the seed
coat and separation of the cotyledons.
Indian subcontinent and the newer high-throughput mills
achieve a dhal yield close to the theoretical maximum of
8385%. Dhal is the main form of consumption in the Indian
subcontinent; the prepared dish, also called dhal, is almost
always part of the main meal. The third main use of chickpea is
as besan, the flour milled from dhal. (Small kabuli seeds have
recently been substituted to produce a cheaper, but inferior
product.) Besan is used for a range of purposes, most com-
monly as a batter or to produce fried, extrusion products.
Chickpea also plays an important, if secondary, role in ani-
mal nutrition. In Mexico, desi types are grown specifically for
inclusion in pig rations. Elsewhere, residues from processing
(seed coat and kibble from milling and damaged or undersized
seeds from grading) are routinely used in intensive livestock
production. Residues from harvest (stems, leaves, pods, and
seeds) are also a valuable feed source for ruminant animals.
Genetic Improvement
Worldwide, there has only been a limited breeding effort in
chickpea compared to the major cereals and oilseeds. Primitive
landraces still account for much of the crop area in developing
programs and the major international centers ICRISAT and
ICARDA. Major breakthroughs have been achieved in develop-
ing resistance to the major diseases ascochyta blight and
fusarium wilt. Cold tolerance, coupled with ascochyta resis-
tance, has been deployed in new varieties that enable winter
sowing in traditional spring-sowing areas, and early maturing
varieties, providing some escape from drought, have facilitated
the wider adoption of chickpeas in central and southern India.
Mechanization of harvest has been facilitated by the develop-
ment of erect plant types, which grow up to 1 m tall and have
stronger, more lodging-resistant branches than the traditional
semierect forms.
There is a paucity of genes for many economically impor-
tant traits in C. arietinum, presumably as a consequence of the
genetic bottleneck caused by domestication. However, genes
conferring improved resistance to many biotic stresses, such as
bruchids, cyst nematode, and phytophthora root rot, can be
found in either or both the closely related wild species
C. reticulatum and C. echinospermum. Breeding programs are
regularly using backcross programs to incorporate a range of
attributes from these species into adapted backgrounds. Intro-
gression of genes from unrelated taxa has also been achieved
by Agrobacterium-mediated transformation. Transformed
plants containing genes with the potential to confer improved
protein quality and insect and disease resistance have been
developed to undergo field evaluation.
In recent years, there have been major advances in chickpea
genomic resources; saturated genetic maps and the publication
of genome sequences of kabuli and desi chickpea types are key
steps that will allow researchers and breeders to develop a full
understanding of the variations found in each genotype.
Exercises for Revision
Why are chickpeas and other grain legumes important for
sustainable farming systems?
What are the three main groups of the species and how do
their end uses differ?
 THE LEGUMES AND PSEUDOCEREALS | Chickpea: Overview
What is the most important disease of chickpea worldwide?
What are the main abiotic constraints of chickpea?
What is the average protein content of the grain?
Which two wild relatives of chickpea are being exploited by
breeding programs?
Exercises for Readers to Explore the Topic Further
The Cicer genus contains nine annual species. How diverse
are they and what is their potential for use in chickpea
improvement programs?
What are the products made from besan (chickpea flour) in
the subcontinent?
See also: Agronomy of Grain Growing: Chickpea: Agronomy;
Appendix 1: Nutrient-Composition Tables for Grains and for Grain-
Based Products; Appendix 2: Lists of Standardized Testing Methods
for the Analysis of Grain and Grain-Based Foods; Food Grains and
the Consumer: Consumer Trends in Grain Consumption; Cultural
Differences in Processing and Consumption; Fortification of Grain-
Based Foods; Genetically Modified Grains and the Consumer; Grains
and Health; Food Grains and Well-being: Functional Foods:
Overview; Food-quality Testing: The Application of Sensory Science
to the Evaluation of Grain-Based Foods; Grain Composition and
Analysis: Standardized Test Methods for Grains and Grain-Based
Products; The Composition of Food Grains and Grain-Based Products;
Units of Grain Science and Trade: Equivalence between the US,
Chinese, and Metric Units; Grains Around the World: Grain
Production and Consumption: Europe; Grain Production and
Consumption in Oceania: Australia and Pacific Countries; Grain
Production and Consumption: Overview; Grain Production and
Consumption: South America; Grain Production and Consumption:
Africa; Production and Consumption of Grains: India; Non-food
Products from Grains: Cereal Grains as Animal Feed; Proteins:
Nitrogen in Grain Production Systems; Nitrogen Metabolism; Protein
Synthesis and Deposition; The Protein Chemistry of Dicotyledonous
Grains; Scientific Organizations Related to Grains: Research
Organizations of the World: Asia-Pacific, CentralSouth America, and
AfricaMiddle East; Research Organizations of the World: CGIAR; The
323
Basics: Taxonomic Classification of Grain Species; The Grain Chain:
The Route from Genes to Grain-Based Products; The Grain Crops: An
Overview; The Legumes and Pseudocereals: Grain Legumes and
Their Dietary Impact: Overview.
Further Reading
Gaur PM, Jukanti AK, and Varshney RK (2012) Impact of genomic technologies on
chickpea breeding strategies. Agron 2: 199221.
Knight R (ed.) (2000) Linking Research and Marketing Opportunities for Pulses in the
21st Century. Dordrecht: Kluwer Academic.
Loss S, Brandon N, and Siddique KHM (eds.) (1998) The Chickpea Book: A Technical
Guide to Chickpea Production. Western Australia: Department of Agriculture and
Food.
Maiti R and Wesche-Ebeling P (eds.) (2001) Advances in Chickpea Science. Enfield:
Science Publishers.
Saxena MC and Singh KB (eds.) (1987) The Chickpea. Wallingford: CAB International.
Saxena NP (1984) Chickpea. In: Goldsworthy PR and Fisher NM (eds.) The Physiology
of Tropical Field Crops, pp. 419452. Chichester: Wiley.
Sharma RN, Shrivastava GK, Rathore AL, Sharma ML, and Khan MA (eds.) (2003)
Chickpea Research for the Millennium. Raipur: Indira Gandhi Agricultural
University.
Singh KB and Saxena MC (eds.) (1999) The Tropical Agriculturalist: Chickpeas.
London: MacMillan.
Smithson JB, Thompson JA, and Summerfield RJ (1985) Chickpea (Cicer arietinum L.).
In: Summerfield RJ and Roberts EH (eds.) Grain Legume Crops, pp. 312390.
London: Collins.
van der Maesen LJG (1972) Cicer L, A Monograph of the Genus, with Special Reference
to the Chickpea (Cicer arietinum L.), Its Ecology and Cultivation. Wageningen: H
Veenman and Zonen NV.
Van Rheenen HA and Saxena MC (eds.) (1990) Chickpea in the Nineties. Patancheru:
ICRISAT.
Varshney RK, Song C, Saxena RK, et al. (2013) Draft genome sequence of chickpea
(Cicer arietinum) provides a resource for trait improvement. Nature Biotechnology
31: 240246.
Yadav SS, Redden R, Chen W, and Sharma B (eds.) (2007) Chickpea Breeding and
Management. Wallingford: CAB International.
Relevant Websites
http://www.ars.usda.gov R&D undertaken on chickpea in the USA.
http://www.cgiar.org Consultative Group on International Agricultural Research.
http://www.icarda.org Information on kabuli chickpeas, mainly in Mediterranean-type
environments.
http://www.icrisat.org Information on chickpeas (mainly desi) in semiarid areas.
 Pea: Overview
TN Khan, University of Western Australia, Crawley, WA, Australia
A Meldrum, Pulse Australia, Perth, WA, Australia
JS Croser, University of Western Australia, Crawley, WA, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Pea is the second most important grain legume crop in the
world and is widely adapted to cooler climates. The major
pea producers are Canada, parts of Europe, the United
States, and Australia.
Pea is used as human food, both as a grain crop and as a
popular vegetable. It is also used for animal feed. Pea seed is
high in protein and a good source of vitamins such as
riboflavin and niacin.
Pea is a climbing plant with a weak stem where tendrils
allow it to trail if support is available. Over the last few
decades, the afila gene that modifies leaflets into tendrils
has been exploited to develop pea varieties with improved
standing ability.
The genetics of the pea plant have been studied in great
detail and dense linkage maps are now available.
Biotechnology techniques including genetic transforma-
tion, embryogenesis, and fast-track technologies are well
established for peas.
Pea production and consumption are likely to remain sta-
ble with increases likely to be proportional to human pop-
ulation increases.
Learning Objective
To understand pea as a grain legume crop in human food
and animal feed and its significance as a legume in crop
rotations.
Introduction
This overview is focussed on the production of the grain
legume crop Pisum sativum (L.), referred to as dry pea or field
pea. Pea is one of the most ancient crop plants domesticated at
around the same time as the major cereals. It is a cool-season
plant requiring moderate mean temperatures in the range of
1218
C with a relatively humid climate for optimum growth.
Hot, dry weather is detrimental to seed set. However, it is also
one of the most adaptable cool-season legumes and can give
economic yields in areas that may not conform to the optimum
requirement. A symbiotic relationship with Rhizobium enables
pea to fix atmospheric nitrogen and make it available for the
following crop. Thus, peas are of considerable significance in
cropping rotations and overall economic value in the cropping
system. The field pea plant has many uses. Green peas are one
of the most popular vegetable items throughout the world.
Some variants with edible pods are also used as a whole-pod
vegetable, for example, sugar-podded peas and snow peas. It is
also a forage plant that makes excellent hay and silage and can
324 Encyclopedia of Food Grains, Second Edition
be used as a green manure crop. Dry peas are used whole in
confectionery and snacks or milled to produce split peas for
making soups, dal (a curried soup-like preparation widely
used in the Indian subcontinent), flour, and canned products
such as mushy peas. Dry peas are also used in the feed industry,
particularly in the diet of pigs and poultry due to their amino
acid balance.
World pea production has declined by 34% since 1993 to
just under 100 million tonnes (Figure 1). Competition for
land from cereals and oilseed crops, which usually deliver
higher profits, is the main reason. There are over 6 million
hectares (Mha) of field pea grown in the world today (Table 1).
This figure has been stable for the last 10 years. As of early
2010s, Canada has the largest area under pea cultivation, fol-
lowed in order by the Russian Federation, China, India, and
France (Table 1). Canada has maintained its pea cultivation
area at over 1.0 Mha since the 1990s, reaching a high of
1.58 Mha in 2008. Canada leads total production with more
than 3.5 million tonnes produced in 2008 and 3.3 million
tonnes (Mt) of crop produced in 2012, followed by the Russian
Federation with a production of 1.6 Mt. Sixty-nine percent of
the total world production comes from the five highest field
pea-producing nations (Table 1).
The yield per hectare varies widely between growing regions
(Table 1). Among the largest producers, France has the highest
yields per hectare of over 4 t against the world average of 1.62 t,
while the leading producer Canadas average yield is  2 t.
Yields generally reflect the degree of industrialization of field
pea production and the inherent variability in the climate.
Average yields in India are below 1 t ha1. In most countries
where it is produced, pea seed is retained for domestic con-
sumption, as are other pulse crops. The United States and
Australia are the other major exporters of field pea.
The Plant
Field pea belongs to the family Fabaceae and although a num-
ber of Pisum species were designated for various cultivated pea
types, they have all been assigned to P. sativum L. Some taxon-
omists, however, still insist on subdividing field pea, garden
pea, and vegetable pea into separate subspecies, although dif-
ferences are often based on a few genes, rather than broad
genomic differences. All subspecies cross and recombine freely.
Despite archaeological evidence indicating its cultivation as
early as 6000 BC in Near Eastern and Greek settlements, there
is no clear consensus on its exact place of origin. The Near East,
Central Asia including Afghanistan, the Mediterranean, and
Ethiopia abound with primitive forms and are sources of
immense genetic diversity. A distinct species Pisum fulvum
Sibth. and Sm., partially cross-fertile with P. sativum, has been
http://dx.doi.org/10.1016/B978-0-12-394437-5.00037-1
 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview 325

Global dry pea production

(1993 to 2013)
16

14

Production (106 tonnes)

12

10

0
1992 1994 1996 1998 2000 2002 2004 2006 2008 2010 2012 2014
Source: FAO Stats

Figure 1 Global dry pea area, production, and yield from 2003 to 2012. Source: FAO Statistic.

Table 1 Area, production, and yield figures for the five highest producers as compared with the world total

Canada Russian Federation China India France World

Aa Pb Yc A P Y A P Y A P Y A P Y A P Y

2002 0.98 1.28 1.31 0.76 1.27 1.67 0.95 1.50 1.58 0.67 0.61 0.91 0.34 1.66 4.92 6.02 9.63 1.60
2003 1.15 1.93 1.68 0.72 1.05 1.47 0.94 1.40 1.49 0.66 0.59 0.89 0.37 1.62 4.41 6.15 9.89 1.61
2004 1.24 3.10 2.49 0.72 1.24 1.72 0.88 1.06 1.21 0.71 0.73 1.02 0.36 1.68 4.71 6.34 11.74 1.85
2005 1.27 2.99 2.36 0.71 1.13 1.58 1.00 1.16 1.16 0.79 0.79 0.99 0.32 1.33 4.21 6.57 11.29 1.72
2006 1.23 2.52 2.05 0.71 1.15 1.62 0.90 1.01 1.13 0.78 0.71 0.91 0.24 1.02 4.21 6.39 9.82 1.54
2007 1.44 2.93 2.03 0.62 0.86 1.39 0.98 1.08 1.10 0.81 0.85 1.05 0.16 0.59 3.61 6.32 9.37 1.48
2008 1.58 3.57 2.26 0.64 1.26 1.98 0.93 1.10 1.19 0.68 0.75 1.10 0.10 0.45 4.50 6.11 10.07 1.65
2009 1.49 3.38 2.27 0.77 1.35 1.75 0.88 0.96 1.10 0.72 0.66 0.92 0.12 0.55 4.72 6.38 10.46 1.64
2010 1.39 3.02 2.17 0.82 1.19 1.45 0.87 0.91 1.05 0.76 0.67 0.88 0.24 1.07 4.44 6.58 10.31 1.57
2011 0.91 2.12 2.31 1.11 2.02 1.82 0.87 1.19 1.36 0.73 0.59 0.82 0.19 0.67 3.58 6.14 9.73 1.58
2012 1.31 3.34 2.14 1.16 1.66 1.43 0.93 1.11 1.20 0.74 0.63 0.85 0.14 0.57 4.05 6.33 9.86 1.56
a
Area: ha  106.
b
Production: t  106.
c
Yield: t ha1.
http://apps.fao.org (February 2015).

identified as a source of useful variation for disease and insect
pest resistance and drought tolerance.
The plant, a spreading or tendril-climbing herb, is typically
30150 cm tall (Figure 2). Papilionaceous flowers may be
white or purple or various shades of purple including pink. It
is predominantly self-pollinated with cross-pollination rarely
exceeding 1%. The stem is typically weak, leading to lodging as
the plant gains biomass. The compound leaves generally have
one or more leaflets modified into tendrils, although variants
are known that have no tendrils. Of significance are the forms
where an entire leaf modifies to form tendrils (Figure 3) as this
may help the crop to stand better by intertwining with plants.
Greater attention to this character will be given while discuss-
ing genetic improvement. White-flowered plants give rise to
white/cream or translucent testae, whereas colored flowers
produce variously colored testae. Cotyledon color ranges
from various shades of yellow to green, although genotypes
with red cotyledon color are also known. The testa color largely
determines the seed appearance, which is of commercial sig-
nificance. Traders recognize five field pea seed types (Figure 4).
The round white (also known as yellow), as the name suggests,
is round with white to cream testae and yellow cotyledons. The
dun type is variously dimpled with greenish brown (dun)
testae and yellow cotyledons. The maple type is round with a
brown testa that is mottled with light-colored spots giving it a
marbling effect and yellow cotyledons. The blue type (also
known as green) has a translucent testa with green cotyledons
giving the seed a bluish hue. The marrowfat is typically a large
seed (>280 mg), slightly dimpled and flattened, with blue/
green testae and green cotyledons.
The diploid chromosome number is 14. Chromosomal
interchanges can be common. Considerable attention has
been given to the genetics of pea following the pioneering
experiments of Mendel in the nineteenth century. Several
326 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview
thousand mutants have been identified and those conforming
to Mendelian genetics were placed on a linkage map as early as
the middle of the last century. A book assembling a compre-
hensive account of pea genome was published toward the end
of the last century. Since then, rapid advances have been made
and are covered in the section Biotechnology and In vitro
Culture Techniques for Field Pea Improvement. Updated
maps, now including biochemical and molecular markers,
have regularly appeared in Pisum Genetics.
Figure 2 P. sativum normal leaf, white-flowered type showing pod and
seed development.
Figure 3 P. sativum types: (a) semileafless, colored flowered type, and (b) normal leaf, white-flowered type.
Nutritional Value, Processing, and Utilization
The field pea seed is a rich source of protein, carbohydrate, and
some minerals, although the nutritional content of the seed
varies with the environment and genetic factors. By far, the
highest proportion of nutrient value in field pea is in the coty-
ledons, with the embryo and seed coat contributing less than
10% to nutritional value. The protein content is typically about
22% but ranges widely, depending on the genotype and growing
conditions (Tables 2 and 3). Although sulfur-containing amino
acid is low, it compares favorably with other grain legumes.
About 60% of the carbohydrate content of the seed is made up
of sucrose and oligosaccharides, starch, and crude fiber. The fat
content is low and the seeds are a good source of vitamins such
as thiamine, riboflavin, and niacin, although considerable loss
of vitamins may occur in processing.
Antinutritional factors are present in quantities lower than
in other grain legumes; the field pea has the lowest trypsin
inhibitor activity among grain legumes commonly used as
food in India. The flatulence activity resulting from oligosac-
charides of the raffinose family is also relatively low, as are
phytic acid and saponins. Lipoxygenase activity that causes
deteriorative changes during the processing and storage of
field peas is high, but these enzymes can be inactivated via
heat treatment.
The dry pea seed is processed via soaking, germination,
milling, cooking, roasting, or fermentation. In Western diets,
the seed is mainly used in soups or in mushy pea preparations.
In South Asian diets, it has varied uses: as dal (spicy soup-like
preparation in the Indian subcontinent), whole pea boiled as a
snack food, as sweet and savory snacks made from pea flour, or
as a supplement to wheat flour to make nutritious breads. In
addition, peas can be processed into protein, starch, and fiber
fractions. These products can then be used in baked goods,
 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview 327

Figure 4 The five seed types of P. sativum: top left, yellow; top right, marrowfat; center, dun; bottom left, green; and bottom right, maple.

Table 2 Nutritional content of field pea seed measured from a Table 3 Amino acid content of field pea seed measured from a
representative number of samples representative number of samples

Nutritional factor Content (g kg1) Amino acid Content (% in seed)

Dry matter 889.0918.0 Alanine 0.97

Protein 193.4273.0 Arginine 2.35
Ash 21.033.1 Aspartic acid 2.46
Fat 7.323.6 Cystine 0.34
Fiber 45.379.0 Glutamic acid 3.85
Acid detergent fiber 76.0176.0 Glycine 0.97
Neutral detergent fiber 71.0245.0 Histidine 0.59
Lignin 0.510.0 Isoleucine 0.93
Calcium 0.51.1 Leucine 1.56
Magnesium 1.01.5 Lysine 1.61
Phosphorus 2.68.5 Methionine 0.19
Potassium 2.011.0 Phenylalanine 0.99
Sodium <0.10.10 Proline 0.97
Sulfur 1.62.2 Serine 1.03
Threonine 0.79
Data from Petterson DS, Sipsas S, and Mackintosh JB (1997) The Chemical Tryptophan 0.18
Composition and Nutritive Value of Australian Pulses. Canberra, Australia: Grains Tyrosine 0.73
Research and Development Corporation. Valine 1.02
Cys Met 0.57
baking mixes, soup mixes, processed meats, health foods, Tyr Phe 1.70
pastas, and purees. There are also several industrial starch uses.
Data from Petterson DS, Sipsas S, and Mackintosh JB (1997) The Chemical
Field peas also serve as excellent animal feed. In an amino
Composition and Nutritive Value of Australian Pulses. Canberra, Australia: Grains
acid-balanced diet, there is no limit on the inclusion rate of Research and Development Corporation.
field peas in rations for sows, weaners, and porkers. For poul-
try, up to 25% can be included in rations. It may also be
fed in high doses to ruminants as a palatable energy and environmental conditions leading to variable yield stability.
protein-rich feed. The logical yield components are pods per plant, seed number,
and seed weight. Among these, pods per plant appear to be the
best correlated with yield. However, selection for yield based
Genetic Improvement on any one of these components is often ineffective. Further,
yield selection in early generations is not very effective, and
Field pea yield is predominantly determined by the additive these generations are used for culling the population against
gene action, but nonadditive factors may also play a significant easily recognized undesirable traits based on flower color, seed
role. The heritability of yield may vary depending on type, disease susceptibility, and plant type. Various breeding
328 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview
methods are in use, and their application depends on the type
and amount of genetic variation in crosses, objectives of
breeding, and available resources. In Europe, the pedigree
method is commonly used. Single-seed descent (SSD) is
becoming popular to hasten breeding and is sometimes used
in conjunction with recurrent selection to increase the fre-
quency of desirable alleles in the population. Bulk pedigree
method has been successfully applied in the Western Austra-
lian short-season environment where biomass appears to be an
important asset in combating terminal drought. Here, bulking
early generations allows for natural selection to favor vigorous
and tall plants.
There is considerable interest in restructuring the field pea
plant since the discovery of a recessive mutant designated as
afila (af) where leaflets are modified into tendrils. Plants with
the afila characteristics are popularly known as semileafless,
although strictly speaking, they should be termed leafless. An
increased number of tendrils in the semileafless crop help the
plant to stand upright through intertwining with adjacent
plants. When combined with dwarf plant stature and stiff
stem characteristics, semileaflessness has considerably
improved the standing ability of the crop. Improved standing
ability has important implications in allowing for efficient
machine harvestability and enhanced aeration through the
canopy that may help to reduce disease epidemics. Semileafless
field pea varieties have become popular in Europe, North
America, New Zealand, and Australia and more recently in
countries such as India. However, in limiting environments,
conventional leaf types still outperform semileafless types. In
such environments, high biomass appears to be an important
factor in determining yield, and semileafless types tend to have
a lower biomass even at a high plant density. As a result,
semileafless cultivars adapted in Australia are much taller and
more vigorous than those in Europe and North America. There
is interest in Australia in conventional leaf type forage field pea
varieties, such as PBA Hayman, that have high biomass pro-
duction and are targeted for the dairy industry.
Breeding against yield limitations such as diseases, insect
pests, and abiotic factors has played a significant role in improv-
ing field pea yields. The impact of these factors varies with
agroecology and crop management practices. Among the dis-
eases, root rot diseases caused by Pythium spp. and Aphanomyces
euteiches f. sp. pisi Pfender and Hagedorn and wilt caused by
Fusarium solani (Mart.) Sacc. f. sp. pisi (Jones) Snyd. and Hans.
are significant. The significant aboveground diseases are (1) wilt
caused by the bacterium Pseudomonas syringae pv. pisi Sackett,
powdery mildew (Erysiphe pisi Syd.), downy mildew (Peron-
ospora pisi Syd.), and black spot disease complex caused by
Ascochyta pisi Lib., Phoma medicaginis var. pinodella (Jones)
Boerema, and Didymella pinodes (Berk. and Blox.) (formerly
known as Mycosphaerella pinodes) Petrak (Figure 5). A number
of viruses attack field peas with the most devastating being pea
seed-borne mosaic virus (PSbMV) and pea enation mosaic virus
(PEMV). Of these, PSbMV is widely distributed throughout the
world, whereas the distribution of PEMV is more restricted.
Resistance is available against some root rots and wilt,
powdery mildew, downy mildew, and black spots caused by
A. pisi and P. medicaginis var. pinodella, as well as the two
important viruses PSbMV and PEMV. However, it is resis-
tance against black spot caused by D. pinodes, the single most
Figure 5 Black spot on P. sativum caused by Didymella pinodes, the
most widespread pea disease in the temperate and Mediterranean
climates.
important disease of field pea throughout the temperate grow-
ing zones, that has been difficult to control through breeding.
Only low to moderate resistance has been identified in several
screening attempts of large germplasm collections in North
America, Europe, and Australasia. More recently, over 3000
accessions from the Vavilov Institute, St. Petersburg, collection
were screened offshore in Ethiopia by a Western Australia-
based project. The more pronounced expression of partial
resistance has been identified in primitive accessions originat-
ing from Afghanistan, Ethiopia, and China. However, such
accessions generally carry a number of wild characteristics
including extraordinarily delayed flowering. Delayed flowering
is recognized to help the field pea plant to resist D. pinodes, and
hence, it is not always clear whether the resistance shown has a
true genetic basis or is an artifact of late flowering habit. There
is some variation in reaction to D. pinodes infection within the
improved germplasm and that a small degree of resistance
scattered in a wide variety of sources can be recombined.
Both wild and improved sources have been used in recurrent
selection programs across Australia and New Zealand, and
some new improved lines show resistance levels that are
equal to or better than the wild germplasm. Availability of
such moderately resistant improved varieties may allow inte-
gration of disease management packages for more effective
disease control.
The major field pea crop pests worldwide are pea and bean
weevil (Sitona lineatus L.), pea moth (Cydia nigricana F.), pea
aphid (Acyrthosiphon pisum Harris), pod borers (Helicoverpa
armigera Hub. and H. punctigera (Wallengren)), and pea seed
weevil (Bruchus pisorum L.). The red-legged earth mite (Haloty-
deus destructor Tucker) is more destructive in the Southern
Hemisphere, for example, Australia, where pasture legumes
are widely grown. There has been some work on studying
variation in resistance levels, but the most promising work so
far has been on pea weevil (B. pisorum) in Australia. Resistance
has been identified in a wild species P. fulvum that is partially
cross-fertile with P. sativum. Resistance has been backcrossed to
field pea lines, and such lines will have application as parents
in crossing.
Waterlogging, freezing temperatures particularly during
flowering and podding, and moisture stress are the most

serious abiotic constraints to field pea productivity. No true
resistance to waterlogging or frost has been reported so far.
There are reports regarding cold tolerance, but whether it
relates to avoiding damage during frost is not known. There
are two aspects of freezing tolerance. In higher latitudes, such
as Europe, freezing tolerance is important during the vegetative
phase in the winter-sown crop, whereas in subtropical and
Mediterranean climates, frost tolerance during the reproduc-
tive stage is more important. A deeper root system has been
reported to increase tolerance to moisture stress. However,
osmoregulation/osmotic adjustment is a more measurable
character, and it has shown good correlation with yield under
water stress in studies done in Spain and Western Australia.
Studies in Western Australia have shown that the most adapted
field pea variety showed one of the highest levels of osmotic
adjustment.
Biotechnology and In Vitro Culture Techniques
for Field Pea Improvement
Molecular Markers and Linkage Mapping
Pea was the model species used by Austrian monk Gregor Men-
del in 1866 to determine the laws of inheritance in plant genet-
ics. Unfortunately, the large size of the pea genome, coupled
with its repetitive nature, has constrained the achievement of a
complete genomic sequence. However, comprehensive genomic
and postgenomic resources have been developed for peas
including BAC libraries, transcriptome and proteome data sets,
and several types of molecular marker sets. Molecular markers
offer the potential to advance plant breeding by analysis of
genetic variation, unequivocal identification of genotypes and
true hybrids, purity testing of breeding lines, genome mapping,
and marker-assisted selection (MAS). In field peas, classical
physiological, biochemical, and hybridization-based markers
have been joined by polymerase chain reaction-generated
markers. The polymorphic markers identified using these tech-
niques have been used to construct dense linkage maps of the
field pea genome, indicating the position and relative genetic
distances between markers along chromosomes.
The recent availability of full genome sequences for three
other legume species has provided opportunities for genome-
wide comparison. This approach has led to the identification
of genes underlying agronomically important traits, some of
which have been applied to MAS programs to improve Pisum
germplasm. When markers that are closely linked to desirable
genes have been identified and validated in specific environ-
ments, plant breeders can use them to select plants on the basis
of genotype, rather than phenotype. The application of MAS
can reduce unreliable phenotypic screening and cultivar devel-
opment time as markers enable the selection of genotypes at
the seedling stage. Molecular linkage maps and trait mapping
are valuable tools for characterizing the genetics of disease
resistance, localizing resistance loci on linkage maps, and iden-
tifying linked polymorphic DNA sequences that might be used
for MAS. This is particularly true for traits that are inherited in a
quantitative manner, for which quantitative trait loci (QTLs)
may be identified. Despite progress in the area of marker
discovery, complete translation to pea breeding is yet to be
achieved.
THE LEGUMES AND PSEUDOCEREALS | Pea: Overview 329
Genetic relationships among Pisum accessions have also
been assessed using genomic techniques. Molecular analysis
of pea collections has shown that although substantial varia-
tion is present within the cultivated Pisum gene pool, wild
Pisum species offer the possibility to incorporate novel traits
that may have been inadvertently eliminated from the culti-
vated gene pool via selection. Association mapping analysis of
diverse pea germplasm will result in the identification of genetic
variation related to desirable agronomic traits. The availability
of high-throughput omics methodologies will translate in the
future into the development of novel, highly accurate selective
breeding tools. Interspecific hybridization to transfer useful
traits from P. fulvum and wild-type P. sativum accessions such
as ssp. P. elatius to cultivated P. sativum would also benefit from
techniques such as MAS and following QTLs. Targeted applica-
tion of existing molecular techniques and the advent of new
techniques such as the analysis of gene function through func-
tional genomics will yield rapid gains in the understanding of
field pea genetics.
In Vitro Mediated Interspecific and Intergeneric Hybridization
Targeted utilization of wild relatives for adaptation to climate
change will almost certainly be an urgent priority this century.
Legume crops and pastures are estimated to use 3560% less
fossil energy than N-fertilized cereal or grasslands. The reduced
energy use was primarily attributed to reduced N fertilizer use
in the legume and subsequent nonlegume crops. Interspecific
hybridization is the crossing of two species from the same
genus. This allows the exploitation of useful genes from wild,
unimproved species for the benefit of the cultivated species.
The Pisum species are diploid self-pollinators sharing a similar
karyotype. P. fulvum is the only separate wild species and a
useful source of disease and insect resistance. Fertile hybrids of
P. sativum  P. fulvum have been produced via conventional
unidirectional crossing with P. fulvum as the pollen donor.
Intergeneric crossing is another technique for improving
genetic diversity within species. It is possible to regenerate
plants from protoplasts in field pea via organogenesis and
somatic embryogenesis. Somatic protoplast fusion, whereby
genomes from different genera are combined without pollina-
tion through protoplast fusion, has been attempted for inter-
generic hybridization between field pea and grass pea. A low
frequency (5%) of protoplast fusion has been achieved; how-
ever, these hybrids have not been regenerated to whole plants.
Transformation
Plant genetic engineering provides an opportunity to introduce
traits such as pest and disease resistance, improved protein
quality, and herbicide tolerance from previously unavailable
sources. Since the first production of diploid transgenic field
pea plants in the early 1990s, Agrobacterium-mediated genetic
transformation has become a routine procedure in a number of
laboratories worldwide. The embryonic axis of immature seeds
or the lateral cotyledonary meristems from germinating seeds
are the most responsive target explants, and regeneration is
generally via organogenesis.
In addition to Agrobacterium-mediated transformation, sev-
eral other approaches have been developed to produce
330 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview
transgenic field pea plants: transgenic callus has been obtained
by electroporation of protoplasts, as has transient expression
following particle bombardment of meristems. Microinjection,
particle gun, agroinfection, and the pollen tube pathway are at
present less efficient or require expensive and sophisticated
equipment. As in other species, transformation efficiency is
dependent on the genotype, explant type, and other physical
parameters. Despite transformation being a well-established
technique in peas, there are currently no commercially avail-
able pea cultivars developed using this technology.
Double Haploid Production
Doubled haploids (DHs) are plants derived from a single imma-
ture pollen grain and doubled artificially to form homozygous
diploids. A DH individual has two identical homologues so that
the amount of recombination information is equivalent to a
backcross. The advantage of using a DH population in molecular
mapping is that all individuals are homozygous. Therefore, DHs
may be transferred between different laboratories and environ-
ments for assessing the effect of the environment on gene expres-
sion. Pea, along with many of the large-seeded leguminous
species, has traditionally been considered recalcitrant to this
technique. In 2009, the first production of confirmed haploid
plantlets of pea from the culture of immature pollen was
reported. Research is ongoing to develop a robust protocol that
will be applicable across a range of genotypes. The mechanisms
for switching the immature pollen from normal gametic devel-
opment have also been recently elucidated for peas. Successful
production of DHs on a routine basis would reduce cultivar
development time and provide excellent recombinant inbred
lines for molecular mapping applications.
In vitro Generation Acceleration
In the absence of a robust, widely applicable, DH protocol, an
in vitro modified SSD system has been developed to accelerate the
achievement of homozygosity in peas. Conventional SSD con-
sists of randomly selecting one seed per plant from an F2 popu-
lation following a cross and advancing it through the early
segregating generations, thus permitting rapid fixation of genes
in breeding lines. As only one seed is needed to produce the next
generation, plants can be grown under conditions that accelerate
flowering and seed set, but do not encourage high yield. Current
SSD methodologies enable a maximum of three generations per
year. The in vitro SSD technique involves further shortening of
the generation time by culturing immature seeds developed after
forced in vitro flowering under conditions for accelerated flower-
ing and seed set. This leads to rapid generation turnover by
reducing the length of the generation cycle, enabling up to
eight generations per year.
Grain Quality and Marketing
Market requirements for grain quality relate predominantly to
shape, size, and color (testa and cotyledon). For all seed types,
large size is desired, and in marrowfat, there may be a limit for
the minimum seed size from the buyer.
With blue peas, resistance to bleaching is an important
criterion. Round white types must have a smooth round sur-
face with white to creamy color and bright yellow cotyledons.
The dun types should ideally have a mix of greenish to brown-
ish testae and fewer and shallower dimples with bright yellow
cotyledons. Maple type, mainly used for pigeon and bird feed,
must have white markings on a brown testa and yellow
cotyledon.
Both round white and dun types are milled, as are blue
types. However, currently, there are no standards for milling
recovery. Seed moisture content at sale is normally between
10% and 15%. All types of field peas are generally accepted by
the feed industry, but a preference is made for the noncolored
type as they tend to have less antinutritional factors. The
Southeast Asian market for sprouting field pea prefers dun
types with large seed size and fewer hard seeds. Irrespective of
the type, the following qualities are important for visual assess-
ment in the trade:
Levels of admixture
Levels of insect damage and presence of live or dead insects
Product color
Product size
Product cleanliness
Product uniformity
Today, almost 50% of the field pea production is traded
throughout the world compared with just 20% 15 years ago
and double that traded in 2002 (Figures 3 and 4), which
compares with temperate cereals where  5% is traded. The
world field pea production has been mostly stable in the last
decade, but trading across the board is showing a strong
increasing trend. The prices have been generally steady since
the early 2000s, but wide variations occur depending upon the
season, time of the year, and proximity to the market. Canada
and the Russian Federation are currently the worlds leading
exporters with Canada having dominated the export market for
many years. The quantity exported from France has halved
since 1999 (Figures 6 and 7). India and China are now the
largest importer of field pea with Bangladesh and Pakistan
importing dry pea as demand requires. Germany, Spain, and
Belgium continue to be consistent importers among European
countries (Figures 8 and 9).
In general, the market for field pea is rather unsophisticated
in terms of both product definition and terminology used. For
instance, in dun-type field peas that are normally milled,
traders often use slight variation in testa color in an attempt
to discount price that is irrelevant to the product utilization.
There are various terminologies used to describe the same
product as in the case of white round type that is traded as
yellow pea from North America. Work continues at an inter-
national level to improve the marketing of pulses through
development of standardized nomenclature and methods of
testing quality parameters in order to meet consumer needs.
The International Pulse Trade and Industries Confederation
(CICILS/IPTIC) and various major importing organizations
are collaborating to develop standardized nomenclature and
methods for quality evaluation that will facilitate improvement
of quality assurance in the international pulse trade including
field peas. Progress is available on the CICILS website (http://
www.cicilsiptic.org/).
 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview 331

Dry Pea Exports by Country 2002

(tonnes)

391,022
Canada
Russian Federation
835,857 672,731
United States of America
France
Australia

Global total (top 20)= 2,703,233

Source: FAO Stats 94,622 130,867

Figure 6 Dry pea exports by the top six exporting countries in 2002.

Dry Pea exports by Country 2011

(tonnes)

243,791
324,883
335,570
Canada
Russian Federation
460,797 2,854,580
United States of America
France
Australia

Source: FAO Stats Global total (top 20)= 4,761,297

Figure 7 Dry pea exports by the top six exporting countries in 2011.

Dry Pea Imports by Country 2002

(tonnes)
India
37,755 99,634
China
91,203
Bangladesh

215,494 Spain
869,803 Belgium
214,545
Pakistan

277,199 Germany

Italy

Global total (top 20)= 2,326,734

110,857
Figure 8 Dry pea imports by the top eight importing countries in 2002.

Future Prospects
The pea crop is likely to retain its importance as human food and
animal feed. While yield improvement per se will continue, seed
quality will become critical as market sophistication develops.
While pea breeding will continue to make improvements in
yield and quality, some serious challenges need to be addressed,
which include both biotic and abiotic stresses.
Among biotic stresses, the black spot diseases and major
insect pests continue to be major limitations to achieving high
yields. Abiotic stresses such as tolerance to drought, frost, and
waterlogging also need attention. Advances in new
332 THE LEGUMES AND PSEUDOCEREALS | Pea: Overview
Dry Pea Imports by Country 2011
(tonnes)
97,345
104,431 102,533 82,527
123,113
211,784
730,434 1,866,735
Source: FAO Stats
Figure 9 Dry pea imports by the top eight importing countries in 2011.
technologies such as those that accelerate generation turnover,
DH, and molecular markers will greatly assist in addressing
future pea breeding objectives. Genetic modification will play a
more dramatic role in addressing some key challenges where
natural variation in pea germplasm is deficient.
Exercises for Revision
What is the status of pea as a crop on the world scene?
What are the climatic conditions most suitable for pea
growing?
What are the uses of pea and how do they relate to seed
characteristics?
What are the major nutritional benefits of pea seeds?
What other advantage does pea crop offer to a cropping
system?
What are the major diseases and insect pests of pea crop?
What is semileafless character in pea plant? Describe the
significance of this characteristic for improving pea varieties.
How does molecular marker technology help in pea
breeding?
Is there a protocol for reliable production of transgenic peas?
Can interbreeding with wild relatives improve pea?
What improvement would you like to see in trading pea
seeds?
Exercises for Readers to Explore the Topic Further
Greater improvement in standing ability of mature pea crop
through combining afila characteristic and greater stem
strength characteristics. What features contribute to stem
strength and their inheritance?
Availability of suitable broad fungal resistance genes for
improving resistance to Didymella pinodes through genetic
modification. Would they be more effective if transferred to
the moderately resistant varieties that have become avail-
able in recent years?
Search for more effective fungicides for D. pinodes control.
Can current fungicides be more effective if used on moder-
ately resistant varieties available now?
India
China
Bangladesh
Spain
Belgium
Pakistan
Germany
Italy
Global total (top 20)= 3,870,336
Continuous improvement of doubled haploid production
to enable reliable and cost-effective techniques for acceler-
ating breeding.
More targeted studies in the available gene pool of P. fulvum
to find traits of interest for transfer to P. sativum.
Frost resistance/tolerance at pod development.
Challenges of possible climate change: heat and drought
resistance/tolerance.
See also: Agronomy of Grain Growing: Field Pea: Agronomy;
Plants: Diseases and Pests; Bioactives and Toxins: The
Antinutritional Components of Grains; Food Grains and the
Consumer: Grains and Health; Genetics of Grains: Development of
Genetically Modified Grains; Genome Mapping; Genomics of Food
Grains; Grain Composition and Analysis: The Composition of
Food Grains and Grain-Based Products; Grains Around the World:
Grain Production and Consumption: China and South-East Asia; Grain
Production and Consumption: Europe; Grain Production and
Consumption in Oceania: Australia and Pacific Countries; Grain
Production and Consumption: Overview; Grain Production and
Consumption: South America; Grain Production and Consumption:
Africa; Production and Consumption of Grains: India; Proteins:
Nitrogen in Grain Production Systems; Scientific Organizations
Related to Grains: Research Organizations of the World: CGIAR;
The Basics: Taxonomic Classification of Grain Species; The Grain
Chain: The Route from Genes to Grain-Based Products; The Grain
Crops: An Overview; The Legumes and Pseudocereals: Grain
Legumes and Their Dietary Impact: Overview.
Further Reading
Byrne OM, Hardie DC, Khan TN, Speijers J, and Yan G (2008) Genetic analysis of pod
and seed resistance to pea weevil in a Pisum sativum  P. fulvum interspecific
cross. Australian Journal of Agricultural Research 59: 854862.
Casey R and Davies DR (eds.) (1993) Peas: Genetics, Molecular Biology and -
Biotechnology. Oxon, UK: CAB International.
FAOSTAT database. Area, yield and production data. http://faostat.fao.org/site/339/
default.aspx; Export and import data. http://faostat.fao.org/site/342/default.aspx
(both accessed 13.12.13).
Hebblethwaite PD, Heath MC, and Dawkins TCK (eds.) (1986) The Pea Crop: A Basis for
Improvement. London: Butterworths.

Khan TN, Timmerman-Vaughn GM, Rubiales D, et al. (2013) Didymella pinodes and its
management in field pea: Challenges and opportunities. Field Crops Research
148: 6177.
Kraft JM and Pfleger FL (eds.) (2001) Compendium of Pea Diseases and Pests, 2nd
edn. St Paul, MN: The American Phytopathological Association.
Makasheva RK (1986) The Pea. Rotterdam: A.A. Balkema.
Ochatt S, Pech C, Grewal R, Conreux C, Lulsdorf M, and Jacas L (2009) Abiotic stress
enhances androgenesis from isolated microspores of some legume species
(Fabaceae). Journal of Plant Physiology 166: 13141328.
Ribalta FM, Croser JS, Erskine W, Finnegan PM, Lulsdorf MM, and Ochatt SJ (2013)
Antigibberellin-induced reduction of internode length favors in vitro flowering and
seed-set in different pea genotypes. Biologia Plantarum 58(1): 3946. http://dx.doi.
org/10.1007/s10535-013-0379-0.
Ribalta FM, Croser JS, and Ochatt SJ (2012) Flow cytometry enables identification of
sporophytic eliciting stress treatments in gametic cells. Journal of Plant Physiology
169: 104110.
Schroeder HE, Schotz AH, Wardley-Richardson T, Spencer D, and Higgins TJV (1993)
Transformation and regeneration of two cultivars of pea (Pisum sativum L.). Plant
Physiology 101: 751757.
Siddique KHM, Erskine W, Hobson K, et al. (2013) Cool-season grain legume
improvement in Australia: Use of genetic resources. Crop and Pasture Science
64: 347360.
THE LEGUMES AND PSEUDOCEREALS | Pea: Overview 333
Smykal P, Aubert G, Burstin J, et al. (2012) Pea (Pisum sativum L.) in the genomic era.
Agronomy 2: 74115.
Relevant Websites
http://www.cicilsiptic.org/ CICILS/IPTIC world trade association for pulses.
http://www.grdc.com.au/ Australian Grains Research and Development Corporation
(GRDC) field pea growing guidelines including disease information.
http://hermes.bionet.nsc.ru/pg/ Pisum Genetics, published periodically by Pisum
Genetics Association, Department of Plant Sciences and Plant Pathology, Montana
State University, Bozeman, MT 59717, USA.
http://www.pea-lentil.com/ USA dry pea and lentil council site with USDA policy
positions and research updates.
http://www.pgro.org/ Processors and Growers Research Organisation (PGRO), United
Kingdom.
http://www.pulseaus.com.au/ Pulse Australia is the peak industry body representing
all sectors of the Australia Pulse Industry.
http://www.pulsecanada.com/ Pulse Canada is the national industry association that
represents growers, processors and traders of pulse crops in Canada.
http://www.saskpulse.com/ Saskatchewan Pulse Growers Group with agronomic
advice, markets and food uses.
 Peanuts: Overview
RCN Rachaputi, University of Queensland, Kingaroy, QLD, Australia
G Wright, Peanut Company of Australia, Kingaroy, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Background, distribution, utilization, and trade.
An overview of morphology, growth, and development.
Yield and quality constraints.
Peanut as a source of oil and quality snack food.
Food safety issues.
Advances in breeding.
Innovations in biotechnology.
Future prospects for increased yield and quality.
Learning Objective
To achieve understanding of the significant issues related to
production, utilization, genetic improvement, and food
safety issues of peanut or groundnut.
Introduction
Peanut or groundnut (Arachis hypogaea L.) is a tropical legume
grown as an oilseed or food crop in more than 100 countries
between 40
N and 40
S, with the main commercial produc-
tion occurring in the northern hemisphere. Peanut area has
increased from 21 million hectares in 2009 to 23 million
hectares in 2013, with an annual production increasing from
33 to 37 million tonnes in the same period.
Peanut kernel is the most economically important compo-
nent with an oil content of 50% and protein content of 25%.
The vegetative parts of groundnut are used as hay for livestock
in developing countries, while the by-products of oil
extraction, that is, shells, peanut meal, and residues, are used
either as livestock feed or as fertilizer in crop husbandry. Pea-
nut being a leguminous plant, can fix atmospheric N (up to
300 kg N ha1), and crop residues incorporated into soil can
potentially contribute up to 120 kg N ha1.
This article provides information on distribution, utili-
zation, and quality issues, especially, on the unique features
of peanut, which lead to its reputation as a high-value crop.
The article also provides an overview of morphology and
recent innovations in biotechnology. Further detail on other
topics not covered in the article can be found in the list of
suggested further reading and related web links.
Historical Background
Peanut is known by many other local names such as earthnut,
groundnut, goober, monkey-nut, pygmy nut, and pignut.
Despite its name and appearance, the peanut is not a nut, but
rather a legume. Peanut genus Arachis, originated in central
334 Encyclopedia of Food Grains, Second Edition
Brazil in South America. The Spaniards are believed to have
taken the crop to the western Pacific, Indonesia, and China
early in the sixteenth century.
Arachis hypogaea is an annual member of the family Papilio-
naceae, which shows a unique feature of maturing their fruits
underground (Figure 1). The plant comprises a number of
subspecies Arachis fastigiata var. fastigiata (Valencia) and
var. vulgaris (Spanish), both bunch forms, and subspecies
Arachis hypogaea var. hypogaea (Virginia bunch and runner).
Large variations in growth habit (erect vs. spreading), branch-
ing type (alternate or sequential), maturity (90160 days), and
traditional market types (Virginia, Spanish, or Valencia) exist.
World Peanut Production and Utilization
Production and Yields
The global nut-in-shell production has averaged around 36
million tonnes during the period 200912, which represents
a 24% increase over 19962002 production figures. Most of
the production occurs in Asia followed by Africa and America
(Table 1). This increase in production has been due to the
combination of increased area planted (nearly fourfold) and
increased yields. Although the average yields have more than
doubled during 19902005, further yield increases have been
modest in most countries during 200912.
Utilization
About 48% of the world production is used as food and 52% is
crushed for oil, although large country variations exist. For
example, India crushes nearly 90% and China 52% of the
production for oil compared with countries such as Indonesia,
Mexico, and Ghana that directly consume over 95% of the crop
for food purposes. In developed countries, about half the
production is used for peanut butter, with the remainder
used for peanut snacks.
Peanut is also an important source of vegetable protein in
addition to 30 essential nutrients. The use of peanut kernels
and their value-added products such as fresh, dry, boiled, or
roasted peanut butter and snack food is on the rise due to
growing vegetarianism and demand for healthy foods. More
recently, peanut has been identified as a source of resveratrol,
an antioxidant linked with reduced cardiovascular disease and
reduced cancer risk.
The peanut haulms are also of very high nutritional value as
livestock feed in developing countries and attract high market
price, particularly in dry years. The shells of the mature pods
can also be used as fuel, fiber filler in animal feeds, garden
mulch, and more recently a possible source for deactivated
carbon (biochar).
http://dx.doi.org/10.1016/B978-0-12-394437-5.00038-3
 THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview 335

Leaf

Flower

Pod

Kernel in pod
Peg

Embryo
Roots

Figure 1 Diagram of a peanut plant showing its unique underground fruiting habit. Source: http://upload.wikimedia.org/wikipedia/commons/a/a4/
Arachis_hypogaea_-_K%C3%B6hler%E2%80%93s_Medizinal-Pflanzen-163.jpg.

Table 1 Annual peanut production and yields for the ten major world-producing countries during the periods

Production % Change
(million Mt) Production (million since Average yield (dry pods) Average yield (dry pods) %Change
Country 200809a Mt) 201112b 200809 (kg ha1) 200809 (kg ha1) 201112 since 2009

China 14.30 16.05 12.2 3.36 3.5 4.2

India 6.25 5.50 12.0 0.92 1.04 13
Nigeria 1.55 2.96 91.0 1.24 1.26 1.6
The United 2.34 1.66 29.1 3.83 3.8 0.8
States
Myanmar 1.00 1.39 39.0 1.49 1.58 6.0
Indonesia 1.25 1.17 6.4 1.67 1.66 0.6
Argentina 0.58 0.98 69.0 3.80 3.19 16.1
Sudan 0.85 0.85 0.0 0.85 0.85 0
Senegal 0.71 0.53 25.4 0.97 0.61 37.1
Vietnam 0.50 0.47 6.0 2.11 2.13 0.9
World 34.43 37.87 10 1.64 1.64 0
production
a
2009 (adapted from www.en.wikipedia.org/wiki/peanut).
b
201112 (adapted from USDA).

Trade
Peanut is generally traded in two product qualities, that is, as
food quality (snack food) and crush quality (cooking oil). The
crush-quality trade is generally mostly domestic to meet
demand for cooking oil. Food-quality peanut in general is
international and receive a quality premium relative to crush-
quality groundnuts.
The world trade market for peanuts can be considered as a
residual market, as only < 6% of the world production is
traded internationally owing to most of the production being
utilized domestically. The major producersexporters of pea-
nuts are the United States, Argentina, Sudan, Senegal, and
Brazil, which in total account for 71% of total world exports.
The peanut trade has steadily shifted from a crush for oil to a
snack food market, which meant rising demand for higher-
quality peanuts, especially with low aflatoxin contamination
(see section Aflatoxin).
More than 48 peanut-growing countries have established
limits for total aflatoxins in food, but standards varied widely,
ranging from 0 to 50 parts per billion. Very strict aflatoxin
regulatory limits in European markets (e.g., <4 ppb total afla-
toxins) have impeded a number of Asian (China, Vietnam, and
India) and African (Nigeria and Sudan) countries in competing
in the edible peanut market.
336 THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview
Morphology and Growth Physiology
Peanut varieties grown in the world can be grouped into four
major botanical types, Spanish, Virginia bunch, Virginia
runner, and Valencia, distinguishable by their branching,
flowering, and fruiting habit.
Being an indeterminate in growth, peanut supports simul-
taneous growth of vegetative and reproductive structures that
compete for nutrients and assimilates throughout most of the
crop life cycle. Flowers appear about 2535 days after planting,
depending on the botanical type of the variety and ambient
temperature during the crop growth, and continue to be pro-
duced until final maturity. The rate of flower initiation
increases to a maximum and then declines as the pod load
increases and the crop approaches maturity. Bursts of flowering
are common when crop recovers from dry spells after rain
during the vegetative phase and can lead to more synchronous
pod set and higher yields and quality given favorable growing
conditions during the pod-filling period. Peanut has a self-
fertilized flower that fades and withers soon after fertilization.
Rates of natural outcrossing are low and have been estimated
between 0.7% and 2.5%.
After 57 days of fertilization, the base of the fertilized ovary
(gynophore or peg) begins to elongate toward the soil, pene-
trating the soil up to a depth of 45 cm. The ovary then begins
to swell and turns horizontally away from the plant (Figure 1),
which takes between 15 and 20 days from the initial flowering
event. There may be 14 kernels per pod depending on the
botanical type. The pod volume expands as the kernels inside
the pod inside fill, reaching full dimensions characteristic of the
variety over the next few weeks. Because of the slow rate of pod
addition, many pods must be added and seeds initiated to form
a full pod load on the plant. Pod growth rates vary depending
on temperature, water status, and varietal maturity and parti-
tioning ability.
Farming Systems
Peanuts are grown as a rotation crop in a range of irrigated or
rainfed cropping systems around the world. In most developed
countries, peanut is grown as a sole crop in rotation with cereal
or sugarcane crops under rainfed or irrigated conditions, using
fully mechanized cultural practices from sowing to harvest. In
countries such as the United States and Australia, it is not
uncommon for one farming family to farm 500 or more hect-
ares per annum. Peanuts are generally grown in a rotation of
one peanut crop in two or three summer cereal crops in order
to minimize soilborne diseases and use of nitrogen fertilizers
and allow more effective weed control.
In developing countries throughout Asia and Africa,
smaller-scale peanut production on 12 ha plots is more com-
mon, under a variety of cropping systems.
In most regions in Asia, peanut is often grown as a cash crop
with both kernels (as food) and haulms (as fodder) are sold in
markets. Peanuts are important component in rice-based crop-
ping systems throughout Southeast Asia. The crop is either
planted with irrigation before a rice crop or grown on residual
soil moisture after the rice harvest. In Asia, peanut is also
grown as an intercrop with longer season annual and perennial
crops, including castor, cassava, maize, and sugarcane. Peanut
can also be grown underneath perennial tree canopies such as
coconut, cocoa, oil palm, and rubber, owing to its effective
shade tolerance. Under these systems, peanut can provide cash
income until the plantation trees are old enough to produce
their end products.
Biotic and Abiotic Constraints to Yield
Biotic Constraints
Insect pests
Around 360 species of arthropod pests are known to attack in
different regions of the world. In general, the foliage pests are
of lesser importance than the soil-inhabiting pests, which can
cause direct economic damage to pods. A large number of
thrips species that attack peanut and have become vectors of
some of the debilitating peanut viruses such as tomato spotted
wilt virus (TSWV). The most important soilborne pests in the
developed world are the lesser cornstalk borer, Elasmopalpus
lignosellus, and white grubs, the larvae of which attack pegs and
pods and cause substantial economic damage. Etiella behrii is
endemic to Australia and is known to be only Etiella species
that affects peanuts. The pod damage caused by the soil insects
can lead to aflatoxin contamination in kernels following infec-
tion via the fungus Aspergillus flavus.
Fungal pests
The most important fungi causing economic damage to peanut
crops around the world are the leaf spots (early and late) and
leaf rust. These diseases cause leaf necrosis and can lead to
severe yield losses if not effectively controlled by fungicides.
A number of highly efficient chemical formulations have been
developed over recent years to control both diseases. Genetic
resistance is also available; however, there is a known negative
association between foliar disease resistance and yield
potential.
A number of soilborne diseases can cause significant eco-
nomic losses in peanuts, via seedling and plant death, as well
as via direct pod damage due to pod rots. The most important
diseases include white mold (Sclerotium rolfsii), Cylindrocladium
black rot, and Sclerotinia blight. More detailed coverage of this
topic can be found in the references suggested in the
bibliography.
Viruses
Many viruses infect peanut, and historically, they have not
caused widespread economic losses. This situation has chan-
ged over the past decade, with the arrival of the potyvirus
peanut stripe virus (PStV) in Asia and the United States and
TSWV in the United States. A system of cultural practices have
been implemented to control these diseases, including plant-
ing date and plant and row spacing to control thrips vectors
such as the western flower thrips. Some genetic tolerance has
been identified for TSWV in the United States, but there is no
known genetic resistance to PStV, although a recent collabora-
tive project involving Australian, Indonesian, and Chinese sci-
entists has developed resistant breeding lines using molecular

approaches. More information on virus-impacting peanut can
be found in the website listed under bibliography.
Abiotic Constraints
About 80% of the worlds groundnut production comes from
dryland regions of the semiarid tropics, which are characteri-
zed by poor soil fertility and drought coupled with elevated
seasonal temperatures.
Drought
The major effect of soil and crop water deficits on peanut yield
relates to reduction in canopy development and pod growth
ultimately affecting pod yield. Peanut has a reputation of being
a drought-tolerant crop, which is associated with a number of
important plant traits. Significant genetic variation in these
traits exists and breeding programs are exploiting these traits
to further enhance peanut drought resistance. Readers are
referred to detailed reviews on this subject by Boote and
Ketring, Wright and Nageswara Rao, and Nageswara Rao et al.
Temperature
Peanut is known to grow well under a wide range of tempera-
tures, with daily optima for the rate of development, growth
processes, and yield being around 2530
C. High tempera-
tures in excess of 35
C can reduce leaf area development,
reduce the number of pods, and result in lowered harvest
indices and yields. The interactions of water deficits with tem-
perature cannot be ignored as high temperatures can accentu-
ate negative impacts of water deficit. In fact, the subterranean
fruiting habit of groundnut renders it more sensitive when soil
surface temperature exceeds 40
C even under well-watered
conditions. The threshold temperature for growth and devel-
opmental processes in peanut is listed in Table 2.
Photoperiod
While growth and yield of peanut are known to be linearly
related to total solar radiation receipt, reproductive develop-
ment and quality can be strongly related to photoperiod (day
length) in sensitive cultivars. Peanuts were previously thought
to be day-neutral, with phenology mainly driven by tempera-
ture. Controlled environment and field experiments have since
determined that reproductive efficiency can be modified con-
siderably by varying photoperiod. Studies have found that
longer photoperiods reduced shelling percentage in sensitive
cultivars. More recent data have shown that significant photo-
thermal interactions exist in peanut, with the sensitivity of
Table 2 Temperature limits of various growth and developmental
processes in groundnut
Process Tbase (
C) Topt (
C) Tmax (
C)
Germination 811.6 2936.5 4147
Leaf appearance 10 30
Leaf photosynthesis 2936
Onset of anthesis 11 2933 38
Pollen viability and seed setting 2732 3743
Pod size and pod growth rate 2428
Harvest index 2732 3744
THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview 337
reproductive efficiency increasing at higher temperatures (e.g.,
mean daily temperatures exceeding 26
C).
Soil nutrition
The specific nutritional issues of most importance for peanut
production include calcium (Ca) and biologically fixed nitro-
gen (BNF).
Peanut pods absorb Ca directly from ambient soil solution.
The Ca absorbed by the roots and transported via the transpi-
ration stream (xylem) is unavailable to the pegs and pods of
the peanut plant as these subterranean organs do not transpire.
Low soil water content in the podding zone can decrease Ca
uptake and induce Ca deficiency in peanut leading to empty
pods, referred to as pops. Calcium, usually applied as gypsum
at during early pod set phase, has been widely practiced to
increase Ca supply to pods and yield of peanut under drought
conditions. Large-seeded Virginia types have been shown to be
more sensitive to drought-induced Ca deficiency than smaller-
seeded Spanish types.
Peanuts as other legumes have the unique ability to fix
atmospheric nitrogen by forming a symbiotic relationship
with Bradyrhizobium bacteria in the root nodules. Inoculation
of seeds with effective rhizobia has led to substantial increases
in peanut yield in fields that have not previously grown pea-
nuts. It is generally accepted that once a field has successfully
grown a peanut crop, it should not require future Rhizobium
applications. Estimates of fixed nitrogen as large as
300 kg N ha1 have been reported for well-watered crops,
with reports that BNF can supply over 80% of the plants
total N uptake. Although most of the N in the crop is removed
when the pods are harvested, numerous experiments have
shown that significant N may be left in the soil for uptake by
subsequent crops. A number of studies showed that the N
fixation process is extremely sensitive to soil water deficits.
Peanut Products, Quality, and Food Safety
Peanut Products
Nearly two-thirds of the world production of peanut is crushed
for cooking oil, making it one of the worlds leading oil crops.
Peanut oil is mostly composed of triglycerides of eight fatty
acids. Around 80% of these fatty acids are either oleic acid
(monounsaturated, C18:1) or linoleic acid (polyunsaturated,
C18:2). Generally, these two fatty acids vary inversely. Of the
eight commonly detectable fatty acids in peanut oil, linoleic
acid is the only one that is polyunsaturated. The second double
bond in a linoleic acid molecule renders it far more susceptible
to oxidation. This oxidative rancidity generates off-flavors in
peanut products and hence limits the shelf life of products in
retail outlets. The monounsaturated oil type in peanuts is also
known to be heart-healthy. It was also found that diets includ-
ing peanuts and peanut butter lowered cardiovascular disease
risk by 21%, whereas the low-fat diet decreased the risk by only
12%.
In the late 1980s, researchers at the University of Florida
discovered a (naturally occurring) mutant peanut plant with
over 75% oleic acid and linoleic acid below 5% of fatty acids,
which they subsequently described as high-oleic acid germ-
plasm. This trait has recently been bred into adapted
338 THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview

commercial varieties, which are currently being released in the
United States and Australia. This characteristic will provide
even greater benefits to both product shelf life and human
health. The increased shelf life is best illustrated by the
peroxide value (PV; a measure of shelf life) data shown in
Figure 2, which shows how high-oleic acid peanuts maintain
low PVs (and hence no rancid off-flavors) during an acceler-
ated aging test.
Nutritional Aspects
Peanuts are an excellent source of nutrition for both humans
and animals. Tables 3 and 4 give a proximate nutritive com-
position of peanut kernels and dry peanut hay, respectively.
Clearly, they are a rich source of protein, oil, carbohydrate, and
minerals. Peanuts are also a good source of niacin, folic acid,
phosphorous, vitamin E, and phytosterols.
Bioactives
Peanuts contain all nine amino acids essential to humans and
nine nonessential amino acids. Recent studies have shown that
peanuts also contain many compounds with antioxidant prop-
erties such as tocopherols, p-coumaric acid, and t-resveratrol,
which can be termed as antioxidants.
products. The human impact, in terms of mortality and mor-
bidity, in peanut-growing countries in developing world is
enormous, with estimates of up to 20 000 deaths per year
arising from aflatoxin-related illnesses being made in some
Southeast Asian countries. In developed countries, consumers
are more aware of food safety issues such as aflatoxin and are
increasingly demanding peanut products meet strict regulatory
standards. Current strategies to reduce aflatoxin levels to below
regulatory limits (e.g., 15 ppb in retail products in the United
States and Australia) involve a post-farm gate process of selec-
tive segregation achieved through blanching and color sorting
in shelling plants. In developing countries, aflatoxin regulatory
limits for peanut have been stringently applied and have
recently affected export of peanut and its products.
A number of potential solutions have been proposed to
minimize aflatoxin at its source, the farm. The use of agro-
nomic management methods, including pre- and postharvest
strategies that minimize aflatoxin contamination by the fungi
Aspergillus flavus and Aspergillus parasiticus, has shown to be
effective. A number of peanut varieties have been shown to
have some aflatoxin tolerance, which seems to be associated
with drought avoidance and escape mechanisms. Recent
research is also investigating the potential for using a bio-
control approach, where nontoxigenic strains of the Aspergillus
flavus fungus are applied to the soil and infect kernels and
hence competitively exclude toxigenic strains.

Food Safety Table 3 Nutritional value per 100 g of peanut kernels

Aflatoxin Principle Nutrient value

Aflatoxin is a human carcinogen that contaminates peanuts,
particularly under end-of-season drought and makes them Protein 2830 g
Oil 4849 g
unsafe for human consumption. Aflatoxin production occurs
Carbohydrate 16 g
in peanut kernels particularly under drought conditions conse- Dietary fiber 8.5 g
quent to infection by soilborne fungi, Aspergillus flavus and Cholesterol 0 mg
Aspergillus parasiticus. Aflatoxin production can occur during Vitamins
crop growth, during postharvest-drying phase, or during storage. Folates 240 mg
The aflatoxin problem is a worldwide phenomenon but is Niacin 12 mg
particularly severe in developing countries in Africa and South Riboflavin 0.135 mg
and Southeast Asia, where food safety and security systems are Thiamin 0.640 mg
not well developed to protect consumers against unsafe food Vitamin E 8.33 mg
Electrolytes
Sodium 18 mg
400 Potassium 705 mg
Minerals
Calcium 92 mg
Peroxide value (meq kg1)

300 Copper 1.14 mg

Normal-oleic
Source: USDA National Nutrient database.
200
Table 4 Nutritional value per kg of dry peanut hay

100 Principle Nutrient value of hay (g kg1)

High-oleic
Crude protein 105
Neutral detergent fiber 466
0 Calcium 12
0 100 200 300 400 500 600 700 800 900
Heating time (h) Phosphorus 1.7

Figure 2 Peroxide value over time as a measure of oxidation of Source: Khan MT, Khan NA, Bezabih M, Qureshi MS, and Rahman A. (2013) The
high- and low-oleic acid peanut oils. Reproduced From OKeefe, Wiley, nutritional value of peanut hay (Arachis hypogaea L.) as an alternate forage source for
and Knauft (1993) Journal American Oil Chemists Society, 70, 489492. sheep. Tropical Animal Health and Production, 45(3), 849853.
 THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview
Cadmium
Cadmium (Cd) is a heavy metal, which can accumulate in
some foods and potentially lead to kidney damage if high
levels of intake occur over prolonged periods. Recent research
has shown that peanut can accumulate high Cd levels on acidic
sandy soils, especially where there has been a history of phos-
phatic fertilizer application, which had contained Cd as a by-
product. A number of management practices have been
proposed to minimize Cd accumulation in peanuts, and readers
are referred to the following website for more information.
Anaphylaxis
Anaphylaxis is a sudden, severe, potentially fatal, systemic aller-
gic reaction to peanuts that can involve various areas of the
body (such as the skin, respiratory tract, gastrointestinal tract,
and cardiovascular system). Anaphylactic reactions can be mild
to life-threatening and seem to have an annual incidence of
about 30 per 100 000 persons. A number of the proteins
responsible for peanut allergy have already been identified.
Among major peanut allergens, Ara h 1, Ara h 2, and Ara h 3,
are storage proteins that comprise a substantial proportion of
the total protein in the seed. These proteins that are critical for
cellular function most likely could not be eliminated without
deleterious consequences for plant growth and development.
However, breeding or bioengineering research is underway to
explore possibilities of substituting a nonallergenic for an aller-
genic form of proteins.
Breeding and Biotechnology
There is a general consensus that further genetic enhancement
of crops including peanut for improved adaptation is best
achieved by trait-based breeding strategies. While conventional
empirical breeding for yield improvement has been generally
successful, the rate of genetic gain has been slow and increas-
ingly becoming cost-prohibitive. Recent studies in Australia
lead to the development of speed breeding strategies, which
can potentially half the time taken for the release of a new
variety with desirable agronomic traits.
There is enormous genetic variability in peanut germplasm
for most traits of interest, including host-plant resistance to
disease and insects and along with important agronomic and
quality traits. This germplasm has been collected and pre-
served. The most significant collection is housed at the Inter-
national Crops Research Institute for the Semi-Arid Tropics
(ICRISAT) in India, where some 12 000 accessions are main-
tained, along with more than 70 wild Arachis species.
Marker-Assisted Breeding
The biotechnology revolution occurring in a range of crops has
yet to be realized in peanut. This has been largely due to the
lack of polymorphism for allozymes or restriction fragment
length polymorphism (RFLP), which has restricted the use of
marker technology for improved selection of quantitative traits
such as yield. The peanut genome is also very large, around
2800 Mbp, and is classed as a tetraploid.
Due to the low number of polymorphic markers available
for the cultivated peanut, there has been very little success in
marker-assisted breeding in peanut. Where the markers have
339
been identified for specific traits such as high oleic acid and
nematode resistance, the marker-assisted selection has been
successful in the development and commercial release of
varieties.
The advent of next-generation sequencing (NGS) provides a
cost-effective strategy to identify molecular markers, at whole-
genome level, associated with traits of interest and can expedite
the development of new peanut varieties. The size and com-
plexity of the cultivated peanut genome have previously been
considered to be a barrier to progress in molecular breeding
research; however, as the cost of NGS continues to decrease,
this appears to be no longer the case.
The International Peanut Genome Initiative (IPGI) estab-
lished in 2004 and currently renamed as Advances in Arachis
through Genomics and Biotechnology (AAGB) is aimed to
facilitate the rapid discovery of progress in the development
of genomic resources across the international peanut commu-
nity. It is expected that a high-quality reference genome for the
cultivated peanut will be available by end of 2015.
Several research institutions have been able to transform
peanut with exogenous DNA and generate fertile plants from
transformed tissues. This technology has opened up enormous
possibilities for expanding the genetic diversity of peanut.
Recent applications of this technology include genetic trans-
formation for disease resistance to PStV and TSWV.
Exercises for Revision
What makes peanut unique compared to other grain
legumes?
What are the major peanut-producing countries in the
world?
What is the major food safety issue with peanut?
What parts of peanut produce aflatoxin?
What factors contribute to aflatoxin production in peanut?
How can aflatoxin contamination be minimized?
What are allergens? What implications do they have for
human health?
Exercises for Readers to Explore the Topic Further
Peanut allergens are becoming a major issue in developing
countries only. What could be the possible reasons?
Allergens are some specific proteins in peanut kernel. How
many proteins are associated with allergens in peanut?
What makes molecular breeding of peanut complicated?
See also: Food Grains and the Consumer: Grains and Health;
Food Grains: Intolerance, Allergy and Diseases: Cereal
Allergens; The Basics: The Grain Chain: The Route from Genes to
Grain-Based Products; The Grain Crops: An Overview; The Legumes
and Pseudocereals: Amaranth: Overview; Beans: Overview;
Chickpea: Overview; Grain Legumes and Their Dietary Impact:
Overview; Pea: Overview; The Oilseeds: Oilseeds: Overview;
Soybean: Overview; Sunflower: Overview.
340 THE LEGUMES AND PSEUDOCEREALS | Peanuts: Overview
Further Reading
Ashley JM (1984) Groundnut. In: Goldsworthy PR and Fisher NM (eds.) The Physiology
of Tropical Field Crops, pp. 453494. New York: Wiley.
Bell MJ, Bagnall DJ, and Harch G (1991) Effects of photoperiod on reproductive
development of peanut (Arachis hypogaea L.) in a cool subtropical environment. II.
Temperature interactions. Australian Journal of Agricultural Research
42: 11511161.
Boote KJ and Ketring DL (1990) Peanut. In: Stewart BA and Nielsen OR (eds.) Irrigation
of Agricultural Crops. Agronomy Monograph 30, pp. 625717. Madison, WI:
American Society of Agronomy.
Chu Y, Holbrook CC, Timper P, and Ozias-Akins P (2007) Development of a PCR-based
molecular marker to select for nematode resistance in peanut. Crop Science
47: 841847.
Chu Y, Wu CL, Holbrook CC, Tillman BL, Person G, and Ozias-Akins P (2011) Marker-
assisted selection to pyramid nematode resistance and the high oleic trait in peanut.
Plant Genome 4: 110117. http://dx.doi.org/10.3835/plantgenome 2011.01.0001.
IPGI (2013) The Peanut Genome Project. In: Wilson RF (ed.) International Peanut
Genome Initiative. http://www.peanutbioscience.com/homepage.html.
Isleib TG, Wynne JC, and Nigam SN (1994) Groundnut breeding. In: Smartt JT (ed.) The
Groundnut Crop, pp. 552623. London: Chapman Hall Chapter 14.
Nageswara Rao RC, Sheshshayee MS, Nataraja Karaba N, Rohini S, Rama N,
Kumaraswamy S, Prasad TG, and Udaykumar M (2012) Genetic approaches to
enhance adaptation of groundnut (Arachis hypogaea L.) to abiotic stresses.
In: Tetuja N, et al. (ed.) Improving Crop Resistance to Abiotic Stress, pp. 304360.
Germany: Wiley-Blackwell, Wiley-VCH Verlag GmbH & Co Chapter 14.
Nigam SN (1992) Groundnut A global perspective. In: Proceedings of an International
Workshop, 2529 November, 1991, 548 pp. Patancheru, India: ICRISAT Centre.
Pattee HE and Young CT (1982) Peanut Science and Technology. Yoakum, TX:
American Peanut Research and Education Association.
Reddy PS (1988) Groundnut. New Delhi, India: Publications and Information Division,
Indian Council of Agricultural Research.
Revoredo CL and Fletcher SM (2002) World Peanut Market: An overview of the past 30
years. Research Bulletin No. 437. The Georgia Agricultural Experiment Stations
College of Agricultural and Environmental Sciences, University of Georgia, Athens,
GA, May 2002. (www.ces.uga.edu/ES-pubs/RB437.htm).
Wright GC and Nageswara Rao RC (1994) Groundnut water relations. In: Smartt JT (ed.)
The Groundnut Crop, pp. 281335. London: Chapman Hall, Chapter 9.
Wright GC, Hubick KT, Farquhar GD, and Rao RCN (1993) Genetic and environmental
variation in transpiration efficiency and its correlation with carbon isotope
discrimination and specific leaf area in peanut. In: Ehleringer JR, Farquhar GD, and
Hall AE (eds.) Stable Isotopes and Plant Carbon-Water Relations, pp. 247268. San
Diego, CA: Academic Press.
Relevant Websites
http://agnews.tamu.edu/stories/PATH/aflares.htm Texas A. & M. University, USA.
http://www.ars.usda.gov/is/AR/archive/jan02/peanut0102.htm Aflatoxin Biocontrol.
http://www.ars.usda.gov/is/pr/1999/990712.htm Agricultural Research Service, USDA.
http://www.clw.csiro.au/publications/general2002/grain_legumes_brochure.pdf
Cadmium in Grain Legumes.
http://cropgenebank.sgrp.cgiar.org/index.php/management-mainmenu-433/stogs-
mainmenu-238/groundnut/guidelines/viruses Peanut Viruses.
http://www.dpi.qld.gov.au/fieldcrops/3027.html Managing Aflatoxin.
http://www.foodallergy.org Anaphylaxis in Peanuts.
http://www.peanut-institute.org/ The Peanut Institute.
http://www.peanutsusa.com The American Peanut Council.
http://sacs.cpes.peachnet.edu/nprl/oldver/atoxin.htm National Peanut Research
Lab, USA.
 Quinoa: Overview
SA Valencia-Chamorro, Escuela Politecnica Nacional, Quito, Ecuador
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Quinoa is an ancient crop from the Andean region.
This grain exhibits great adaptability to extreme climate,
from sea level to 4000 m, and is able to grow in poor and
marginal soils with low annual rainfall.
It presents a wide genetic variability with differences in
form, in grain color and size, in grain yield and also in
terms of its by-products.
Quinoa, a very nutritious grain crop, is a unique grain,
which contains the correct balance of essential elements
and vitamins, and it is gluten-free.
The main producers of quinoa are Bolivia, Peru, and Ecua-
dor. However, production is expanding to other continents
such as Africa, Asia, Australia, and Europe.
Quinoa production is around 80 000 t (8  107 kg)
annually.
Processed foods from quinoa include a wide range of bread
types, pasta, noodles, breakfast cereals, beer, porridge, etc.
NASA has designated quinoa as a potential crop for its
Controlled Ecological Life Support System (CELSS).
Quinoa presents high potential to contribute to food secu-
rity and nutrition for humankind in several regions
worldwide.
Learning Objectives
To achieve understanding of the adaptability, variability,
nutritional quality, and uses of quinoa grain, the ancient
crop from the Andean region.
To be able to assess the future prospects for quinoa as a
major grain crop.
Introduction
Quinoa (Chenopodium quinoa, Willd) is an indigenous crop of
the Andean region of South America. It is one of the oldest
crops of the American continent. Archaeological findings in
northern Chile showed that quinoa was used prior to 3000
BCE. In Ayacucho, Peru, evidence has been found that quinoa
was cultivated before 5000 BCE.
The quinoa plant was widely cultivated in the entire
Andean region, in Colombia, Ecuador, Peru, Bolivia, and
Chile, before the Spanish conquest. However, habits and tra-
ditional foods of the native people were replaced by foreign
crops such as wheat and barley. Therefore, quinoa was culti-
vated either in small plots in rural areas for domestic consump-
tion or as borders for other crops such as potatoes or maize. For
that reason, it was classified as food for poor people.
Quinoa is grown in the highlands from 5  N in southern
Colombia to 30  S in northeastern Argentina. At sea level, it is
cultivated between 36  S and 40  S, in central Chile. Thus,
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00041-3
quinoa is cultivated from sea level in Chile to 4000 m in the
Altiplano of Peru and Bolivia.
In the late 1970s, the main production areas of quinoa were
located in the Andean region. In Peru and Bolivia, this crop was
of great importance. It was cultivated not only for domestic
consumption but also for export. The cultivation of quinoa
then spread from the Andean region to several countries in the
world. In the 1980s, quinoa was grown in Canada, Denmark, the
United States, and the United Kingdom. In the late 1990s, an
American and European project on quinoa to promote the
regional interchange of excellent genetic material among research
institutes and universities from both continents was undertaken.
Today, after more than 20 years of work, quinoa has worldwide
distribution. Quinoa is being cultivated in several countries
of the Andean region (Colombia, Bolivia, Ecuador, Chile, and
Peru), in Africa (Kenya), Asia (China, Japan, the plains of
Northern India, and the Himalayas), Australia, Europe (England,
Denmark, France, Germany, Italy, the Netherlands, Portugal,
Spain, and Sweden), and North America (United States and
Canada). Quinoa production is in a process of expansion into
different geographic areas of the planet due to its ability to adapt
to a wide range of agroecological zones with extreme ecological
conditions.
Classification
Quinoa belongs to the Chenopodiaceae family, genus Cheno-
podium. Its botanical name is C. quinoa, Willd. Common names
used in the Andean region are quinua, kiuna, or parca in
Ecuador, Peru, and Bolivia; supha, jopa, jupha, jiura, aara,
ccallapi, or vocali in Bolivia; quinhua in Chile; and suba or
pasca in Colombia.
The classification of quinoa was first made based on the
color of the plant and fruits. Subsequently, it was based on the
morphological types of the plant. Despite the wide variation
observed, quinoa is considered to be one single species. For
practical reasons, quinoa, like maize, was classified in terms of
races.
The most extensive collection of different quinoa races
belongs to Peru and Bolivia; each has over 2000 ecotype sam-
ples. However, other collections exist in Argentina, Colombia,
Chile, Ecuador, England, the United States, and the former
Soviet Union.
Quinoa collected in Ecuador, Peru, and Bolivia has been
classified into 17 races; however, more races may exist. Two
types of inflorescence are described:
1. Glomerulates are small groups of flowers (glomeruli)
originating from tertiary axes.
2. Amaranthiformes have glomeruli originating mainly from
secondary axes.
According to this, the races of quinoa are classified as follows:
first, glomerulate inflorescence (Cajamarca, Copacabana,
341
342 THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview

Cuzco, Challapata, Cochabamba, Sicuani, Junn, Ancash, clustered in different forms either amaranthiforme or glomer-
Glorieta, and Dulce) and, second, amaranthiforme inflo- ulate. The flowers are incomplete; they do not have petals.
rescence (Achacachi, Puno, Real, Potosi, Puca, Sucre, and Quinoa has both hermaphrodite flowers, located at the distal
Pichincha). end of a group, and female flowers located at the proximal end.
Quinoa grows from sea level to the Andean highlands. Thus, The fruit of quinoa is an achene. It is covered by
one of the most useful classifications is that describing five perigonium, which is the same color as the plant: white, yel-
ecotypes (Table 1): sea level, valley, subtropical, salar, and low, gray, light brown, pink, black, or red (Figure 2). It is easily
altiplanic. removed when it is dried. It produces small, circular-shaped
Quinoa presents a great genetic diversity, with differences in seeds, range 1.54 mm (1.5  10 34.0  10 3 m) diameter
color, form, and size, as well as in quality and quantity of (250500 seeds per gram).
primary elements and secondary metabolites. Quinoa germ-
plasm banks in the Andean region have been in existence since
the 1960s. Germplasm collection has continued in several
Andean countries. The main quinoa germplasm collection is
preserved in Bolivia, which conserves more than 5000 quinoa
accessions. Peru, Ecuador, and Chile hold more than 536, 642,
and 25 quinoa accessions, respectively.

Taxonomic and Botanical Description

Quinoa is a plant of the Chenopodiaceae family, genus Che-

nopodium, section Chenopodia, and subsection Cellulata. The
genus Chenopodium is the largest in the family Chenopodia-
ceae and has worldwide distribution, with about 250 species.
With regard to botanical description, quinoa is an annual
gynomonoecious plant; it can grow up to 3 m high, depending
on the variety and the environment (Figure 1).
The seeds can germinate very rapidly, in a few hours after
being exposed to moisture. Roots can reach depths of up to
0.30 m. The stem is cylindrical (0.030.05 m diameter); it can
be either straight or with branches, and its color is variable.
Depending on the variety, it changes from white, yellow, or
light brown to red. The leaves are shaped like a gooses foot.
They are formed by petioles and lamina; petioles are long-
channeled on their upper side. The lamina is polymorphous in
the same plant: rhomboidal or triangular in the lower lamina of
leaves and triangular or lanceolate in upper leaves. Quinoa
inflorescence is a panicle, full of bunches (racemose), which
emerge on the upper part and do not have branches. The length
of the panicle varies from 0.15 to 0.70 m. Flowers can be Figure 1 Quinoa plant. Courtesy of Ing. Angel Murillo, INIAP Ecuador.

Table 1 General ecotypes of quinoa

Growth
Ecotypes Location altitude (m) Varieties Characteristics

Sea level South of Chile <500 Chilean varieties Unbranched, long day plants (<2 m tall),
yellow, bitter seeds
Valley Andean valley 20004000 Blanca de Junn, Rosada de Jun, Big plants (23 m tall), branched, short growth
Amarilla de Matangani, Dulce de period
Quitopamba, Dulce de Lazo
Subtropical Subtropical area of 25003000 Plants with intense green color that turn orange
Bolivia (Yungas) as they mature; small seeds, white or orange
Salar Bolivian Salares 37003800 Real Plants adapted to salty and alkalinic soils;
bitter seeds; high saponin content
Altiplanic Area around Lake 35004000 Cheweca, Kancolla, Blanca de Jul Short plants with straight stems (11.8 m tall);
Titicaca short growth period; resistant to frost

Source: Risi J and Galwey NW (1984) The Chenopodium grains of the Andes: Inca crops for modern agriculture. Advances in Applied Biology 10: 145216. Reproduced from
Encyclopedia of Food Sciences and Nutrition, second ed. (2003), p. 4896. Elsevier Ltd.

Figure 2 Quinoa seeds. Courtesy of Silvia Valencia Chamorro.
Two layers enclose the seed. The pericarp adheres to the seed.
It contains saponins, which confer the bitter taste characteristic
of quinoa. The episperm encloses the seed as a thin layer. The
embryo is formed by two cotyledons and a radicle; it comprise
about 60% of the volume within the episperm and forms a ring
around the perisperm. The high protein content of quinoa,
unlike cereals, is explained by the high proportion of embryo.
Cultivation Practices and Production
The cultivation of quinoa is related to the crop rotation. In the
Andean region, traditional rotation includes maize, faba beans,
potatoes, lupin, wheat, and barley. This practice improves
quinoa yield and preserves soil fertility.
Cultivation of the quinoa plant requires loose soil that can
retain an adequate amount of moisture. However, it can be
cultivated in different areas in South America with a wide range
of rainfall (0.052.00 m). Quinoa tolerates a wide range of pH
conditions of the soil (from pH 4.8 to 8.5), relative humidity
(from 40% to 80%), and temperatures (from 4 to 38  C).
Quinoa is frost-resistant. Therefore, quinoa presents great
adaptability to extreme climatic and soil conditions. For this
reason, quinoa has been ranked among 21 grain species most
resistant to climate change along with beans, maize, amaranth,
onions, and others.
Sowing density may vary according to region. It has
been reported from 0.4 to 0.6 g m 2 (4.0  10 4 to
6.0  10 4 kg m 2) in the Bolivian Altiplano, from 0.5 to
2.3 g m 2 (5.0  10 4 to 2.3  10 3 kg m 2) in Puno, and
from 1.2 to 1.6 g m 2 (1.2  10 3 to 1.6  10 3 kg m 2), for
varieties such as INIAP-Tunkahuan and INIAP-Pata de Venado,
in Ecuador. However, sowing density can depend on varieties
and climatic conditions of the region where it is cultivated.
Quinoa is harvested at physiological maturity. Physiologi-
cal maturity may be reached within 7090 days after flowering.
Depending on the variety, it takes between 5 and 8 months for
a plant to mature.
Traditionally, harvesting is done manually. The plants are
cut with a sickle and then placed in windows to dry completely.
The yield of quinoa can be in the range of 45500 g m 2
THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview 343
(4.5  10 25.0  10 1 kg m 2) depending on the variety and
growing conditions. Quinoa has a seed coat containing sapo-
nins, which confer a bitter taste. The saponins must be
removed before consumption. External layers are removed
using either a wet or a dry process. The traditional wet process
used in rural areas is hand scrubbing in alkaline water. This
process is used on a commercial scale; it involves abrasive
dehulling to remove the external coverings, followed by a
thorough washing. However, this method has economic and
ecological restrictions: The water demand is high and waste-
water is contaminated with saponins, which are toxic to
cold-blooded animals. Moreover, wet grains must be dried
immediately, or they may germinate after a few hours in wet
conditions. In the dry method, grains are scrubbed and
polished to remove, as fine powder, their external layers. This
method presents several advantages; no water is needed, no
heat treatment to dry the seeds is required, and no environ-
mental contamination is produced. This method is best suited
to sweet varieties (low saponin content) of quinoa.
A combination of dry and wet processes is applied to bitter
varieties (with high saponin content) of quinoa. Saponin is
first removed by polishing, when most of the saponin is
removed. Then, the saponins that remain in the seeds are
washed with water, followed by a dry process. Any of the
processes described earlier makes the quinoa ready for use by
the consumer or for further processing. In the future, a novel
enzymatic method for removing saponins utilizing enzymes of
Eurysacca quinoae Povolny may be used. However, this method
has not yet been commercialized.
The most important fungal disease is downy mildew (Per-
onospora farinosa), which requires high humidity and tempera-
ture. The main symptom is chlorotic lesions on the upper
surfaces of the leaves, with a white or purple mycelium on
the lower surfaces. Another disease is brown stalk rot, which
is caused by Phoma exigua var. foveata. The stem often becomes
shrunken and the plant may become chlorotic with progressive
defoliation toward the apex may occur. Kcona kcona (Scrobi-
palpula sp.) is probably the most serious pest of quinoa. Larvae
first destroy leaves and inflorescences. Later, when the plant is
mature, larvae destroy the panicle and grains.
Today, where quinoa is cultivated in new agroecological
regions, it is probable that it will provide a new host for the
local pests and fungi.
The production of quinoa is consistently increasing world-
wide. The main producers are Bolivia, Peru, the United States,
Ecuador, and Canada. In the 6 years, from 2005 to 2011, the
cultivated area of quinoa increased by around 47%, from
68 863 to 101 527 ha (6.9  1081.0  109 m2), and the pro-
duction increased about 37%, 58 443 to 80 241 t (5.8  107
8.0  107 kg). However, quinoa is still mainly cultivated in the
Andean region: Bolivia, Peru, and Ecuador. Bolivia is the main
producer country, with 54% of cultivated area and 51% total
production. Bolivia is also the main exporter of the grain,
followed by Peru and Ecuador. Several efforts have been
made to increase the production of quinoa in the Andean
region, for example, in Ecuador, where the potential cultivated
area for quinoa production in 2012 was 86 856 ha
(8.7  108 m2).
Traditional production of quinoa was based on crop rota-
tion. However, in some Andean areas, quinoa is grown as an
344 THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview

intensive monoculture using mechanization. Thus, some con-
cerns with regard to environmental sustainability have arisen.
Chemical Composition and Nutritional Value
The diet of ancient inhabitants of the Inca Empire has gener-
ated interest due to its being extremely nutritious.
Like cereals, quinoa grains contain carbohydrates, protein, fat,
minerals, fiber, and vitamins (Table 2). The chemical composi-
tion of quinoa depends on the variety, soil, cultural practices, and
environment of its cultivation. Values for the nutritional charac-
teristics of 555 quinoa accessions have been reported: protein
(10.2118.39%), fat (2.0510.88%), dietary fiber (3.469.68%),
ash (2.125.12%), carbohydrates (52.3172.98%), and energy
(1.31  1041.68  104 kJ kg 1). Quinoa accessions show wide
variability for most of the studied parameters, which is indicative
of their genetic potential.
Quinoa Grains
Protein
The protein content of quinoa grains can vary from 8% to 22%,
which on an average is higher than that of common cereals
grains such as rice, maize, and barley. However, it represents
less than half of the quantity of protein in most legumes. In
quinoa, most of the protein is located in the embryo. The
nutritional value of quinoa varies considerably among varie-
ties, which may depend on soil and fertilization regime. Sev-
eral studies on the chemical composition of quinoa have been
made in the past decades. For instance, in Bolivia, variation in
nutritional values of 20 different quinoa accessions from the
Banco Nacional de Germoplasma de Granos Altoandinos
(BNGA) was reported. The protein content was in the range
12.4715.48%, which is low.
Albumins and globulins are the major protein fraction
(4477% of total protein) in quinoa, which is greater than
the prolamins (0.57.0%). Thus, quinoa is considered to be
a gluten-free grain because it contains very little or no prola-
min. Therefore, quinoa provides a nutritional food source,
which is of particular relevance to gluten-intolerant people.
The good nutritional quality of quinoa is primarily due to
its high protein content and quality. Quinoa protein has a
good balance of essential amino acids (Table 3). It is excep-
tionally high in lysine, an amino acid, which is generally
not overly abundant in plant foods. It is also a good comple-
ment for legumes, which are often low in methionine and
cystine.

Table 2 Chemical composition of quinoa and some cereals and legumesa (10 2
kg per kg dry wt)

Quinoa Barley Corn Rice Wheat Oat Rye Bean Lupin Soybean

Protein 16.28 11.02 10.51 7.94 17.65 18.40 11.57 26.72 40.39 39.90
Fat 7.00 1.29 5.29 2.92 2.20 7.52 1.82 0.94 10.88 21.80
Ash 2.74 1.23 1.34 1.53 2.17 1.87 1.76 4.34 3.66 5.32
Carbohydrateb 73.99 86.44 82.85 77.24 77.98 72.21 84.85 68.00 45.08 32.98
Fiber, total dietary 8.07 17.35 8.14 3.50 13.98 11.55 16.89 28.22 21.10 10.28
kJ kg1(104)c 1.54 1.47 1.53 1.55 1.34 1.63 1.41 1.39 1.55 1.87
a
Quinoa, uncooked; Barley, pearl, raw; Corn, yellow; Rice, brown, long-grain, raw; wheat, hard red spring; Beans, kidney, all types, mature seeds, raw; Lupins, mature seeds, raw;
Soybeans, mature seeds, raw.
b
Carbohydrate includes fiber.
c
kJ kg1: 4  (%protein + carbohydrates) + 9  (%fat).
Source: Adapted from US Department of Agriculture, Agricultural Research Service. 2013. USDA National Nutrient Database for Standard Reference, Release 26. Nutrient Data
Laboratory Home Page, http://www.ars.usda.gov/nutrientdata (accessed 21-02-2014).

Table 3 Essential amino acids in quinoa and other foodsa (103 kg per kg protein)

Quinoa Corn Rice Wheat Bean Milk FAO/WHO/UNU

Histidine 29 11 25 21 51 30 15
Isoleucine 36 36 42 35 44 52 30
Leucine 60 70 83 67 80 95 59
Lysine 54 32 38 26 69 84 45
Methionineb 36 29 35 41 26 32 22
Phenylalaninec 61 75 89 77 15 10 38
Threonine 30 31 37 28 42 43 23
Tryptophan 12 11 13 13 12 13 6
Valine 42 49 57 44 96 63 39
a
Quinoa, uncooked; Corn, yellow; Rice, brown, long-grain, raw; Wheat, hard red spring; Beans, kidney, all types, mature seeds, raw; Milk, whole, 3.25% milk fat, with added vitamin D.
b
Methionine + cystine.
c
Phenylalanine + tyrosine.
Source: Adapted from US Department of Agriculture, Agricultural Research Service (2013). USDA National Nutrient Database for Standard Reference, Release 26. Nutrient Data
Laboratory Home Page, http://www.ars.usda.gov/nutrientdata (accessed 21-02-2014); World Health Organization (2002). Joint FAO/WHO/UNU Expert Consultation on Protein and
Amino Acid Requirements in Human Nutrition. WHO technical report series; no. 935. Geneva: WHO.
 THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview 345

The nutritional evaluation of quinoa protein has been Table 4 Mineral composition (105 kg per kg dry wt) and vitamin
reported in several studies. The protein efficiency ratio of raw concentration (105 kg per kg dry wt) of quinoa and some cerealsa
debittered quinoa was 7893% of that of casein. These figures
Quinoa Wheat Rice Barley
increased when quinoa was cooked and became 102105% of
that of casein. Thus, the protein quality of quinoa matches that Minerals (105 kg per kg)
of the milk protein casein. Ca 54.2 28.7 25.7 32.3
Mg 227.2 142.1 159.5 87.9
Carbohydrates and fiber K 649.2 389.7 248.8 311.4
The major component in quinoa is carbohydrates, which varies P 526.9 380.6 371.5 245.8
from 67% to 74% of the dry matter. Starch is about 5260%. Fe 5.3 4.1 1.6 2.8
The starch is located in the perisperm of the seed; starch may be Cu 0.7 0.5 0.3 0.5
Zn 3.6 3.2 2.3 2.4
as simple units or as spherical granules. The diameter of quinoa
Vitamins (105 kg per kg)
starch granules is in the range of 0.42.0 mm (4  10 7
Thiamin (B1) 0.42 0.58 0.45 0.21
2  10 6 m). Starch granules in quinoa are smaller than those Riboflavin (B2) 0.37 0.13 0.10 0.13
reported for maize (range 123 mm (1.0  10 6 Niacin (B3) 1.75 6.55 5.68 5.12
2.3  10 5 m)) and for wheat (240 mm (2  10 6 Ascorbic acid NR 0.00 0.00 0.00
4  10 5 m)). The gelatinization temperature for different qui- -Tocopherol 2.81 1.16 1.34 0.02
noa cultivars was found to be in the range of 55.572.0  C. The -Carotene 0.01 0.01 0.00 0.01
starch amylose content is about 11%, which is lower than in
NR, not reported.
cereals, for example, rice (17%), wheat (22%), and barley a
Quinoa, uncooked; wheat, hard red spring; rice, brown, long-grain, raw; barley, pearl,
(26%). The total dietary fiber ranges from 11.0 to 13.3% in
raw.
raw and heat-treated quinoa. The crude fiber content is in the Source: Adapted from US Department of Agriculture, Agricultural Research Service
range 2.53.9%, and pentosans 2.93.6%. The high content (2013). USDA National Nutrient Database for Standard Reference, Release 26. Nutrient
of dietary fiber has many positive health effects, for example, it Data Laboratory Home Page, http://www.ars.usda.gov/nutrientdata (accessed 21-02-
can reduce the level of cholesterol in the blood and improve 2014).
digestion. For this reason, consumers in developed countries
may also have an interest in including quinoa in their diets.
Other carbohydrates are found in small amounts, such as acid, hederagenin, 30-o-methylspergulagenate, and phytolac-
monosaccharides (2%) and disaccharides (2.3%). cagenic acid are identified in sweet and bitter genotypes of
quinoa. Saponins show hemolytic activity and are in general
Fat, vitamins, and minerals toxic to cold-blooded animals, which obtain oxygen directly
Quinoa contains from 2% to 10% fat. Quinoa and soy oils from water.
exhibit similar fatty acid composition. Thus, quinoa is a rich Reduction of plasma cholesterol has been attributed to the
source of essential fatty acids such as linoleic (18:2n 6: 52%) presence of certain saponins in the diet. However, some sapo-
and linolenic (18:3n 6: 4%). The fatty acid composition is nins can form insoluble complexes with minerals, such as zinc
similar to maize germ oil. Quinoa is also a good source of and iron, which make the minerals unavailable for absorption
minerals (Table 4). It contains more calcium, magnesium, in the gut.
iron, and zinc than common cereals, and the iron content is Phytic acid (myoinositol 1,2,3,4,5,6-hexakis dihydrogen
particularly high. phosphate) is found in most cereals and legumes, fruits, and
Polishing and washing quinoa grain reduce the mineral vegetables. In quinoa seeds, phytic acid is located in the external
content to some extent, by 1215% for iron, zinc, and potas- layers as well as in the endosperm. Even small amounts
sium, 27% for copper, and 3% for magnesium. Quinoa con- (0.5 mmol g 1 (5.0 10 4 mmol kg 1)) of inositol hexapho-
tains more riboflavin (B2) and a-tocopherol than rice, barley, sphate (IP6) or pentaphosphate may reduce the solubility of
or wheat. The concentrations of the natural antioxidants g- and iron. IP6 was mainly found in varieties from Ecuador: sweet
a-tocopherol in quinoa were found to be 797.2 and 788.4 ppm INIAP-Tunkahuan (11.3 mmol g 1 (1.1  10 3 mmol kg 1)) and
(7.97  10 47.88  10 4 kg kg 1), respectively. Thus, quinoa bitter INIAP-Ingapirca (8.6 mmol g 1 (8.6  10 3 mmol kg 1)).
can be a source of vitamin E. These levels were reduced to just 0.3 mmol g 1 (3.0 -
 10 4 mmol kg 1) by fermentation of germinated quinoa
Other components flour. At the same time, a five- to eightfold increase in the amount
Saponins (after removal), phytic acid, trypsin inhibitors, and of soluble iron was found.
tannins are present in quinoa in low levels. As stated, saponin, The trypsin inhibitor activity in eight varieties of quinoa
a bitter-tasting compound, is located in the outer layers of (range 1.365.04 TIU mg 1 (1.36  10 65.04  10 6 TIU -
quinoa seeds (at levels of 0.15%). It protects the seeds from kg 1)) was found to be much lower than in soybean (24.5 -
birds and insects. The amount of saponins present depends on TIU mg 1 (2.5  10 5 TIU kg 1)). Trypsin inhibitor is a
the variety. Bitter-flavor varieties contain more saponin than thermolabile protein, which is inactivated by heat treatments.
sweet varieties. Saponins are glycosidic compounds, which The total phenolic content (using the FolinCiocalteu
occur in two groups. According to the nature of the sapogenin method) of precooked quinoa seeds indicated that bitter seeds
moiety, they are conjugated with hexoses, pentoses, or uronic contained higher amount of phenolics (range 7.2310.05 mg
acids. The sapogenins are steroids (C27) or triterpenoids gallic acid equivalent per 100 g (7.2  10 51.0  10 4 kg gallic
(C30). Using gas chromatography, the sapogenins oleanolic acid equivalent per kilogram)) than sweet ones (range
346 THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview
6.059.39 mg/100 g (6.1  10 59.4 10 5 kg gallic acid equiv-
alent per kilogram)). Moreover, bitter quinoa precooked seeds
exhibited a higher flavonoid content (1.041.74 mg catechin
equivalent per 100 g (1.0  10 51.7  10 5 kg catechin equiv-
alent per kilogram)) than sweet seeds (0.710.91 mg/100 g
(7.1 10 69.1 10 6 kg/kg 1)). Thus, bitter and sweet quinoa
seeds varieties are a rich source of antioxidant compounds, the
former being higher in antioxidants.
Quinoa Leaves
The leaves of quinoa are comparable to spinach as regards
flavor. They contain (on a wet basis) carbohydrate 4.8%, pro-
tein 3.3%, fat 1.8%, ash 3.3%, and fiber 1.9%. The protein
content of quinoa leaves is also similar to spinach. However, its
protein contains slightly more isoleucine (5.8% per 100 g pro-
tein) and valine (7.5% per 100 g protein). The amount of
saturated fatty acids such as palmitic (16:0; 16.7%) and stearic
(18:0; 1.3%) is higher than in the grains. Quinoa leaves are a
rich source of vitamin A containing 2085 mg RE (retinol equiv-
alents) per 100 g (fresh wt.) (2.9  10 5 kg RE per kilogram
(fresh wt.)) and vitamin E 2.9 mg a-TE (alpha-tocopherol) per
100 g (2.9  10 8 kg a-TE (alpha-tocopherol) per kilogram).
Fresh quinoa leaves contain more magnesium (83 mg per
100 g fresh wt. (8.3  10 4 kg per kilogram fresh wt.)) and
sodium (289 mg per 100 g fresh wt. (2.9  10 3 kg per kilog-
ram fresh wt.)) than spinach leaves. Also, quinoa leaves have
low levels of the antinutrients nitrate and oxalate.
Sapogenins in the leaves increased as the plant matures.
After 120 days of from sowing, the sapogenin content in leaves
of sweet genotypes varied between 0.013% and 0.017% (dry
matter) and in bitter varieties varied between 0.02% and
0.16% (dry matter). Hederagenin was the major sapogenin
present in the leaves.
Uses
In Chile, Ecuador, Peru, and Bolivia, the main uses of whole
quinoa seeds are in soups, salads, casseroles, and stew. Cooked
grains swell to about two or three times their original size.
Figure 3 Quinoa products. Courtesy of Lcdo. Guillermo Tapia, Cereales Andinos Ecuador.
Quinoa can be eaten as a rice replacement, as a hot breakfast
cereal, or boiled in water to make an infant cereal (Figure 3).
The grains can even be popped like popcorn. The seeds can be
also sprouted. The sprouts need to turn green before they can
be added to salads.
Quinoa flour can be mixed with maize or wheat flour for
making bread and pasta, and it is used for making biscuits,
cakes, and pastries in general.
Several levels of substitution of quinoa flour have been
reported, for instance, in bread (1013% quinoa flour),
noodles and pasta (3040% quinoa flour), and sweet biscuits
(60% quinoa flour). All levels produce quinoa products of
excellent quality. Quinoa flour can also be drum-dried and
extruded, producing products with good physical, sensorial,
and nutritional qualities. Solid-state fermentation of quinoa
with Rhizopus oligosporus Saito has been performed, to produce
tempeh of good quality.
In Bolivia, in 1975, the government adopted a resolution
mandating that 5% of quinoa flour must be added to all pastas,
crackers, and breads.
Quinoa leaves, like grains, can also be cooked, made into a
spinach-like dish, or may be served raw in a salad.
During four years, from 2002 to 2006, an Andean Quinoa
Project was undertaken in three countries, Bolivia, Peru, and
Colombia. Research goals included agriculture and agro-
industrial activities. Several quinoa enriched quinoa products
were developed. Additionally, more than 30 different pro-
cessed products from quinoa grains and plants have been
reported. Quinoa has jumped from a staple food to gourmet
kitchen food.
Saponins from quinoa, due to their chemical qualities, have
multiple purposes. They are used in the manufacture of soap,
liquid detergents, shampoos, toothpaste, stain removers, and
foam stabilizer; in beverages, especially beer; as an insecticide;
in a compound for fire extinguisher; and in photo processing.
The grains, the stems, and the leaves have been used for their
medicinal properties, for cicatrizing, as an anti-inflammatory, an
analgesic, and a disinfectant. The plant-dried stalks are used as
fuel or may be used in preparing bleach or dyes or for cardboard
and paper. The natural colorant from quinoa (betalains) was
traditionally used for cloth dyeing and food preparation.

Because of its excellent nutritional characteristics, NASA has
included quinoa as a potential crop for its CELSS project. The
CELSS concept utilizes plants to remove carbon dioxide from
the atmosphere and generate food, oxygen, and water for the
crews of long-term human space missions.
Future Perspectives
Quinoa has long been recognized as nutritious food. The
importance that quinoa could play in nutritional behavior is
now being emphasized, not only in developing countries but
also in the developed world. Quinoa could be a strategic crop
used to complement the diet in rural/marginal regions where
protein-energy malnutrition affects most of the population.
On the other hand, quinoa represents an exotic and healthy
rediscovery in the developed world. Germplasm collection
should continue in countries of the Andean region.
The production of the grain has notably increased in the
Andean region. It is also in a process of expansion in other
parts of the world. The potential for improving quinoa pro-
duction seems promising. Therefore, new varieties (precocity,
grain size and color, resistance to biotic and abiotic factors, and
grain yield) and by-products should be obtained through
genetic improvement. Participatory plant breeding for quinoa
(including plant breeders, agronomists, communities, farmers,
and others) is desirable. Improvement of quinoa yields by
improving cultivation practices should be performed.
There is need for intensive cultivation of quinoa. This could
meet the quantity and quality needs of the food industry.
Traditional production of quinoa includes crop rotation. In
the Bolivian Southern Altiplano, production is often made
without crop rotation. This could lead to very unsustainable
production systems. Further research is needed in order to
identify the agroecological and social basis of quinoa sustain-
ability in traditional and new cultivation regions.
Improving methods for removing saponins without signif-
icant modification of nutritive value is encouraged. The selec-
tion of sweet genotypes with low saponin content in the seeds
is the main breeding goal. Further research is needed to find
markers for indirect selection of sweet genotypes.
Aggressive promotion campaigns should be carried out to
encourage greater consumption of quinoa, as it is a very healthy
and gluten-free grain. Quinoa has been spread outside of the
Andean region, owing to its great adaptability to different agro-
ecological systems. The year 2013 was declared the
International Year of the Quinoa. Several international events
were carried out all over the world. Quinoa should be promoted
as a food of the twenty-first century, as it has a high potential to
contribute to food security in several regions worldwide.
Exercises for Revision
What region is the origin of quinoa? Where is most genetic
variation to be found?
What is the adaptability of this crop to soil and climate
conditions? Could quinoa be produced near where live?
THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview 347
What important nutritional characteristics does quinoa
have? How can these be exploited in development of new
food products?
Which environmental concern is related to increased pro-
duction in the Bolivian highlands?
How is quinoa being promoted all over the world? Do you
think it has potential to be major grain worldwide?
Exercises for Readers to Explore the Topic Further
Quinoa is a unique food crop, which contains all the essen-
tial amino acids in the correct balance. Taking this into
account, what are the agro-industrial prospects for this
grain?
Production of quinoa was originally in Andean countries;
now, quinoa is cultivated in other countries. What are the
challenges arising from this fact?
See also: Carbohydrates: Carbohydrate Metabolism; Cereals:
Chemistry and Physicochemistry of Non-starchy Polysaccharides;
Glycemic Index, Glycemic Load, and Their Health Benefits; Grains other
than Cereals: Non-starch Polysaccharides; Health Effects of b-Glucans
Found in Cereals; Resistant Starch and Health; Starch: Analysis of
Quality; Starch: Chemistry; Starch: Starch Architecture and Structure;
Starch: Synthesis; Fats: Healthy Fats and Oils; Lipid Chemistry; Food
Grains: Intolerance, Allergy and Diseases: Celiac Disease;
Cereal Allergens; The Gluten-Free Diet; Grain Composition and
Analysis: The Composition of Food Grains and Grain-Based
Products; Proteins: Nitrogen in Grain Production Systems; Nitrogen
Metabolism; Protein Synthesis and Deposition; Proteomic Analysis
Relevant to Cereal Grains; The Enzymes Associated with Food Grains
and Their Functions in Processing; The Protein Chemistry of
Dicotyledonous Grains; Thiolomics of the Gluten Protein Network of
Wheat Dough; The Basics: Taxonomic Classification of Grain
Species; The Grain Chain: The Route from Genes to Grain-Based
Products; The Grain Crops: An Overview; The Grains that Feed the
World.
Further Reading
Brkic M, Garca A, 2013 Un cultivo ancestral para apuntalar el futuro. http://www.
alimentosargentinos.gov.ar/contenido/revista/pdfs/57/57_04_QUINOA.pdf
(accessed 01.10.13.).
FAO 2011 Quinoa: An ancient crop to contribute to world food security. http://www.fao.
org/fileadmin/templates/aiq2013/res/en/cultivo_quinua_en.pdf (accessed
01.02.13.).
Gandarillas H (1979) Historia y distribucion geografica. In: Tapia M, Gandarillas H,
Alandia S, Cardoso A, Mujica A, Ortiz R, Otazu V, Rea J, Salas B, and Zanabria E
(eds.) La Quinua y la Kaniwa. Cultivos Andinos, pp. 2144. Bogota: Centro
Internacional para el Desarrollo.
Giusti K (1970) El genero Chenopodium en la Argentina. I. Numero de Cromosomas.
Darwiniana 16: 98105.
International Year of the Quinoa. www.fao.org/quinoa-2013 (accessed 13.08.13.).
Jacobsen SE, Mujica A, and Ortiz R (2003) The global potential for quinoa and other
Andean crops. Food Rev. Int. 19: 39148.
Masterbroek H, Limburg H, Gilles T, and Marvin H (2000) Occurrence of sapogenins in
leaves and seeds of quinoa (Chenopodium quinoa Willd). J. Sci. Food Agr.
80: 152156.
Mujica A, Jacobsen SE, Izquierdo J, 1998 Libro de Campo: Prueba Americana y
Europea de Quinua (Chenopodium quinoa Willd). Universidad Nacional del
348 THE LEGUMES AND PSEUDOCEREALS | Quinoa: Overview
Altiplano, Puno, Peru; Red de Cooperacion Tecnica en Producion de Cultivos
Alimenticios de la Oficina Regional de la FAO para America Latina y el Caribe.
Mujica A, Ortiz R, Bonifacio A, et al., 2006. Proyecto Quinua: Cultivo Multiproposito
para los Pases Andinos PNUD- PROY/INT/01/K01- Peru- Bolivia- Colombia.
Informe Final.
Peralta E, Mazon N, Murillo A, et al., 2012. Manual Agrcola de Granos Andinos:
Chocho, Quinoa, Amaranto y Ataco. Tercera edicion. Publicacion Miscelanea No.
69. Programa Nacional de Leguminosas y Granos Andinos. Estacion Experimental
Santa Catalina, INIAP.
Repo-Carrasco R, Espinoza C, and Jacobsen S-E (2003) Nutritional value and use of the
Andean crops quinoa (Chenopodium quinoa) and Kaniwa (Chenopodium
pallidicaule). Food Rev. Int. 19: 179189.
Risi J and Galwey NW (1984) The Chenopodium grains of the Andes: Inca crops for
modern agriculture. Adv. Appl. Biol. 10: 145216.
Soto JL and Carrasco E (2012) Market potential of Andean grains. In: Giuliani A,
Hintermann F, Rojas W, and Padulosi S (eds.) Biodiversity of Andean Grains:
Balancing Market Potential and Sustainable Livelihoods, pp. 2574. Rome, Italy:
Bioversity International.
Tapia M (1979) Historia y distribucion geografica. In: Tapia M, Gandarillas H, Alandia S,
Cardoso A, Mujica A, Ortiz R, Otazu V, Rea J, Salas B, and Zanabria E (eds.)
La Quinua y la Kaniwa. Cultivos Andinos, pp. 1119. Bogota: Centro Internacional
para el Desarrollo.
U.S. Department of Agriculture, Agricultural Research Service, 2013. USDA
National Nutrient Database for Standard Reference, Release 26. Nutrient
Data Laboratory Home Page, http://www.ars.usda.gov/nutrientdata
(accessed 21.02.14.).
Valencia S, Svanberg U, Sandberg A-S, and Ruales J (1999) Processing of quinoa
(Chenopodium quinoa Willd): Effect on the in vitro iron availability and phytate
hydrolysis. Int. J. Food Sci. Nutr. 5: 203221.
World Health Organization, 2002. Joint FAO/WHO/UNU expert consultation on protein
and amino acid requirements in human nutrition. WHO Technical Report Series: No.
935. WHO: Geneva.
 Grains Around the World
Contents
Grain Production and Consumption: Overview
Grain Production and Consumption: Africa
Production and Consumption of Grains: India
Grain Production and Consumption: China and South-East Asia
Grain Production and Consumption: Europe
Grain Production and Consumption: Cereal Grains in North America
Grain Production and Consumption: Oilseeds in North America
Grain Production and Consumption in Oceania: Australia and Pacific Countries
Grain Production and Consumption: South America
Grain Production and Consumption: Overview
T Beta and C Isaak, University of Manitoba, Winnipeg, MB, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Wheat, rice, and maize contribute over 90% of global cereal
production.
World production of wheat, paddy rice, and maize was 652,
701, and 875 million tonnes (Mt) in 2012.
The major wheat-producing regions are the EU, FSU, NC
America, South America, Oceania, and Asia.
The major rice exporters include India, Thailand, Vietnam,
the United States, and Pakistan.
Maize is the preferred crop in Africa and South America.
World oilseed production was 454 Mt in 2012 with soy-
bean as the predominant grain crop.
Learning Objectives
Achieve understanding of the major grains that are pro-
duced and consumed worldwide.
Introduction
The term grain is used to include cereals (wheat, rice, and the
coarse grains such as maize, barley, sorghum, oats, and rye),
oilseeds, and pulses. World grain production, particularly in
wheat and rice, increased substantially between 1961 and 1980
due to the increase in yields per hectare or the Green
Revolution (Figure 1). Since then, per capita (per person)
grain production has not increased much as most of the easily
realizable benefits of plant breeding, fertilizer, machinery,
and irrigation have already been achieved. Production of
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00051-6
grain per hectare is close to the maximum obtainable through
photosynthesis; hence, world grain production has not
matched the increase in worlds population. Low grain prices
at planting time, high temperatures, and water shortages are
the major factors contributing to reduced grain harvests. The
fall in production has triggered an increase in the prices of
wheat and maize and a corresponding increase in products
(bread, breakfast cereals, pasta, and livestock products, includ-
ing meat, milk, and eggs) derived from these grains.
World production of wheat, rice (paddy), and maize in
2012 was 652, 701, and 875 million tonnes (Mt), respectively
(Table 1). Total cereal production is shown in Figure 2 as an
average of 10-year periods beginning 1961. There was a 30%
increase in cereal production during the period between
197180 and 198190. The rate of increase in cereal produc-
tion has been slowing since the days of the Green Revolution in
the 1970s. Table 1 shows world production of cereals, wheat,
rice, maize, barley, sorghum, millet, and rye for the period
201012. Data on yields and area harvested are also listed.
The worlds wheat was produced in Asia (47%), Europe
(29%), and North and Central (NC) America (13%) in 2012.
About 90% of the worlds rice was produced in Asia. Africa and
South America contributed 4% and 3% to the worlds rice
production, respectively. NC America, Asia, South America,
and Europe contributed 36, 33, 13, and 11%, respectively, of
the worlds maize production. Wheat, rice, and maize contrib-
uted to over 90% of global cereal production. World exports
and imports of cereals by region in 2009 and 2010 are given in
Figures 3(a) and 4(a), respectively. Pulses (including dry
beans, dry peas, chick pea, and pigeon pea) were 3% of the
total cereal production. Among oilseeds, soybean was 10% of
world cereal production.
349
350 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview

2800
2600
2400
2200
2000
Production (mt)

1800
1600 Cereals, total (Mt)
1400 Rice, paddy (Mt)
1200
Wheat (Mt)
1000
800 Maize (Mt)
600
400
200
1960
1961
1962
1963
1964
1965
1966
1967
1968
1969
1970
1971
1972
1973
1974
1975
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
Year

Figure 1 World grain production for the period 19612012. FAOSTAT Database 2013.

Table 1 World cereal production for the period 201012 Wheat

Area harvested (Mha) Yield (kg ha 1) Production (Mt) Data on world wheat production, area harvested, and yield for
the period 201012 are given in Table 1. World wheat produc-
Cereals, total
tion decreased by 49 Mt in 2012 from the previous year
2010 693 3571 2547
2011 707 3660 2589 (Table 1). The area harvested declined by 4 million hectares
2012 703 3622 2474 (Mha) and yields were 60 kg ha 1 less than in 2011. The
Wheat average global wheat production increased tremendously dur-
2010 217 3003 675 ing the period 196080 largely due to an increase in yield. In
2011 221 3175 701 developed countries, wheat yields tripled between 1950 and
2012 217 3115 652 1996 due to new wheat varieties and improved technologies
Rice, paddy including methods of sowing, irrigation, fertilizer application,
2010 162 4336 718 moisture retention, and integrated pest management. Potential
2011 163 4429 723
yields are still to be realized in African and Asian developing
2012 163 4395 701
countries due to several constraints including inadequate
Maize
2010 164 5187 850 water, infertile soils, poor weather, and lack of inputs. The
2011 172 5154 885 major wheat-producing countries during 2012 by regions are
2012 177 4944 875 EU (France, Germany, and the United Kingdom), FSU (the
Barley Russian Federation and Ukraine), NC America (Canada),
2010 47 2606 124 South America (Argentina), Oceania (Australia), and Asia
2011 48 2747 133 (China, India, Pakistan, Turkey, Iran, and Kazakhstan).
2012 49 2684 132 Bread wheat is planted on 93% of world wheat-growing
Sorghum area while durum and soft wheat occupy the remainder. Spring
2010 41 1439 59
types are sown on two-thirds of the land devoted to wheat in
2011 42 1384 59
developing countries.
2012 38 1535 58
Millet The five major wheat exporters are France (EU), the United
2010 36 905 32 States and Canada (NC America), the Russian Federation
2011 34 803 27 (FSU), and Australia (Oceania) (Figure 3(b)). India and
2012 31 820 26 Pakistan produce enough wheat to meet their country needs.
Oats Developing countries in Africa, Asia, and South America
2010 9 2163 20 accounted for at least 50% of world wheat imports in 2009
2011 10 2348 23 and 2010 (Figure 4).
2012 10 2179 26 Figure 5 shows consumption of wheat in the EU, FSU, NC
Rye
America, South America, NE Asia, FE Asia, and Africa in 2009.
2010 5 2374 12
Most wheat is used for human consumption with a limited
2011 5 2565 13
2012 6 2617 15 amount going into livestock feed. World wheat production and
consumption are projected to increase in 2013 and 2014.
Source: FAOSTAT Database 2013. Exports to China are expected to increase in 2013 due to a
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview 351

Average cereal production (Mt)

700

600

400

300

100

0
1961-70 1971-80 1981-90 1991-2000 2001-10

Average wheat production (Mt)

700
600
500
400
300
200
100
0
1961-70 1971-80 1981-90 1991-2000 2001-10

Average paddy rice production (Mt)

700
600
500
400
300
200
100
0
1961-70 1971-80 1981-90 1991-2000 2001-10

Average maize production (Mt)

800
700
600
500
400
300
200
100
0
1961-70 1971-80 1981-90 1991-2000 2001-10
Figure 2 World cereal production, averages for 10-year periods. FAOSTAT Database 2013.

poor quality domestic crop. Wheat dry milling produces flour, bran, a by-product of milling, is used in high-fiber foods and as
semolina, bran, and germ. The primary products of milling are an animal feed ingredient. Wheat is also used as a valuable feed
used largely in baking and extrusion to produce breads and ingredient for milk and beef livestock as it is more nutritious
pastas. Wheat germ oil and meal are valuable coproducts. The than maize, sorghum, and barley.
 Other
ME Asia
0% Africa
6% Africa S America Other
ME Asia 1%
FE Asia 1% 10% 1%
FE Asia 7%
6%
6%
S America
12%

EU
25% NC America EU
31% 26% NC America
33%

FSU FSU
15% 8%
Oceania Oceania
(a) 6% 6%

S America
ME Asia Other S America Other
ME Asia 5%
3% 1% 5% 1%
5%
Africa
0% Africa
FE Asia 0%
0% FE Asia
EU
0%
32% EU
NC America
35% NC America
29%
32%

FSU FSU
20% Oceania 11%
(b) 10% Oceania
11%

Other Africa S America Africa

0% 4% 9% Other 3% S America NC America
0% 6% 12%
NC America
10% Oceania
ME Asia ME Asia
19% 0%
Oceania 22%
0% FSU
FSU
0%
0%
EU EU
5% 6%

FE Asia FE Asia
53% 51%
(c)

ME Asia Africa
Other 2% ME Asia Other Africa
3%
FE Asia 3% 2% 2% 2%
2%
FE Asia
1%
EU
EU S America FSU 15%
16% 18% 3% S America
27%
Oceania
FSU 0%
8%
NC America NC America
Oceania
48% 48%
0%

(d)

Figure 3 (a) World exports of cereals by region for 2009 (330 Mt) and 2010 (334 Mt); regional totals exclude processed secondary products.
(b) World exports of wheat by region for 2009 (146 Mt) and 2010 (145 Mt); regional totals exclude processed secondary products. (c) World exports of
milled rice by region for 2009 (30 Mt) and 2010 (33 Mt); regional totals exclude processed secondary products. (d) World exports of maize (corn)
by region for 2009 (101 Mt) and 2010 (108 Mt); regional totals exclude processed secondary products. FAOSTAT Database 2013.
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview 353

Other
Other 3%
4%

Africa Africa
19% ME Asia 20%
ME Asia
18%
20%
S America S America
7% 8%

NC America
NC America FE Asia
FE Asia 8%
8% 23%
21%
Oceanla
EU Oceanla EU
FSU 0%
21% 0% 20%
FSU 0%
(a) 0%

Other
Other
3%
3%

Africa
ME Asia Africa
ME Asia 24%
17% 26%
23%

S America FE Asia
FE Asia 7% 17%
15% S America
NC America 9%
EU 5% EU
23% Oceanla NC America
23%
0% 5%
FSU FSU Oceanla
(b) 0% 0% 0%

Other
Other 4%
4%

Africa Africa
ME Asia 31% ME Asia 29%
23% 24%

S America
FE Asia 4%
S America FE Asia
18% NC America
EU 4% 21% EU
10% NC America 9% 6%
7% Oceanla
FSU Oceanla FSU
(c) 1% 2% 1% 2%

Other Other
2% 3%

Africa Africa
ME Asia ME Asia 13%
14% S America
12% S America 13% 8%
9%

NC America NC America
12% 12%
FE Asia
33% FE Asia
35% Oceanla
EU 0%
EU Oceanla FSU
16%
18% 0% 0%
FSU
(d) 0%

Figure 4 (a) World imports of cereals by region for 2009 (328 Mt) and 2010 (335 Mt); regional totals exclude processed secondary products.
(b) World imports of wheat by region for 2009 (147 Mt) and 2010 (144 Mt); regional totals exclude processed secondary products. (c) World imports of
milled rice by region for 2009 (30 Mt) and 2010 (31 Mt); regional totals exclude processed secondary products. (d) World imports of maize by
region for 2009 (100 Mt) and 2010 (107 Mt); regional totals exclude processed secondary products. FAOSTAT Database 2013.
354 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview

100%

80%

60%

40%

20%

0%

EU

ica

ica

ia

ia

ca

ld
FS

As

As

or
ri
er

er

Af

W
Am

Am

FE
M
NC

S
(a)

100%

80%

60%

40%

20%

0%
EU

ica

ica

ia

ia

ld
ric
FS

As

As

or
er

er

Af

W
Am

Am

FE
M
NC

(b)

100%

80%

60%

40%

20%

0%
EU

ica

ica

ia

ia

ld
ric
FS

As

As

or
er

er

Af

W
Am

Am

FE
M
NC

(c)

Food Feed Food Manufacturing Seed Other

Figure 5 Wheat, maize, and milled rice consumption by regions for 2009 in tonnes. (a) Total wheat consumption in selected regions were 117 (EU 15),
54 (FSU), 46 (NC America), 24 (South America), 170 (ME Asia), 130 (FE Asia), 52 (Africa), and 610 (world). (b) Total rice consumption (milled
equivalent) in selected regions were 63 (EU 15), 5 (FSU), 325 (NC America), 59 (South America), 36 (ME Asia), 208 (FE Asia), 62 (Africa), and 770
(world). (c) Total maize consumption in selected regions were 3 (EU 15), 1 (FSU), 5 (NC America), 15 (South America), 130 (ME Asia), 245
(FE Asia), 21 (Africa), and 423 (world). FAOSTAT Database 2013.

Rice
World production of paddy rice (unmilled or rough rice) was
701 Mt in 2012, a decrease of 22 Mt from the previous year.
The area harvested remained the same but yields decreased by
34 kg ha 1 during the same period. World rice production
doubled from 216 to 448 Mt during the period 196183. The
increase in average rice production was 15% between the
10-year periods 198190 and 19912000 and 19912000
and 200010 compared with 30% for the periods 197180
and 198190 (Figure 2).
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview
Most of the rice is grown in developing countries. In 2012,
Asia was responsible for 93% of the world rice production,
51% of the total crop being produced by China and India.
Only small amounts (< 10%) of world rice produced are
traded internationally. World exports of rice by region for
2009 and 2010 are given in Figure 3(c). The major rice
exporters include India, Thailand, Vietnam, the United States,
and Pakistan. Many countries in NC and South America,
Europe, and Africa are net importers (Figure 4(c)). Total rice
consumption (milled equivalent) for 2009 in Mt was 3
(EU15), 1 (FSU), 5 (NC America), 15 (South America), 130
(ME Asia), 245 (FE Asia), and 21 (Africa), giving a total of 423
(world). Global rice trade is predicted to remain the same in
2013 and 2014 as there are limited opportunities for expan-
sion. Both production and consumption are expected to
increase slightly worldwide.
Coarse Grains
World production of coarse grains (including maize (corn),
barley, sorghum, oats, rye, millet, triticale, buckwheat, and
quinoa) was 1153 Mt in 2012 versus 1121 Mt in 2010. Among
the coarse grains, maize, barley, and sorghum are the most
important in terms of total cereal production. World produc-
tion of maize, barley, and sorghum was 875, 132, and 58 Mt,
respectively, in 2012 (Table 1). Production of other coarse
grains was below 30 Mt each. A record of 1295 Mt, an increase
of 11.4% from 2012 in global production of coarse grain in
2013, has been attributed to a strong recovery in maize pro-
duction in the United States following the 2012 drought-
reduced level. Leading producers in coarse grain production
include the United States, China mainland, EU, Brazil, India,
Argentina, the Russian Federation, Ukraine, Mexico, Canada,
Nigeria, Indonesia, Ethiopia, Turkey, and South Africa. Major
exporters of coarse grains are Argentina, Australia, Brazil,
Canada, EU, Russian Federation, Ukraine and the United
States.
Maize
Maize composes 35% of total cereal production in the world.
Maize is the preferred crop in Africa and South America. In
Asia, it occupies the third position after rice and wheat. World
maize production fell by 10 Mt from 885 Mt in 2011 (Table 1).
The area harvested in 2012 was 177 Mha. Yields have remained
relatively low in developing countries (1980 kg ha 1) com-
pared with the global average (4944 kg ha 1) due to environ-
mental, technological, and socioeconomic factors. Developed
countries make use of adequate inputs and a well-mechanized
system for maize production. In 1961, world maize production
was 205 Mt. The figure rose to 615 Mt in 1998. After the
1998 peak in world maize production, rising production costs
and shortage of foreign exchange in many developing countries
have led to diminished production and, consequently, maize
trade. There was a 50% increase in average global maize pro-
duction between the periods 196170 and 197180 (Figure 2).
World exports and imports of maize by region for 2009 and
2010 are given in Figures 3 and 4, respectively. The main maize
355
exporters are the United States, Argentina, Brazil, France, Hun-
gary, and Ukraine. The United States accounts for almost half
of the world maize exports. China supplies grain to its neigh-
boring countries. Japan and the Republic of Korea are the
largest importers, each importing 16 and 9 Mt, respectively,
in 2010.
Globally, the total domestic supply of maize was 799 Mt in
2009. The primary product of maize milling is the meal used
for feed and food purposes. At least 55% of world maize
production is used to feed livestock and 14% is used for
human food. Other uses include industrial processing and
seed. Secondary and derived products are obtained through
dry-milling and wet-milling processes. Global maize trade is
forecast to reach a 6-year high in 2013 as the demand in China
is increasing.
Barley
World barley production was 132 Mt in 2012, down by 1 Mt
from 2011 (Table 1). The area planted has steadily increased by
1 Mha from 2010 to 2012 but yields declined by 63 kg ha 1
from 2011 to 2012. In 2012, the major producers by region
included the EU (Germany 10, France 11, and Spain 6), FSU
(Russian Federation 14), NC America (Canada 8 and the United
States 5), Asia (Turkey 7), and Oceania (Australia 8). The EU is a
major exporter of barley. The Russian Federation and Ukraine
have become competitors with the EU in export markets.
Exports from the EU also face competition from Australia
and Canada especially for supply of malting barley. ME Asia
and FE Asia imported 10 and 4 Mt, respectively, of the total 25
Mt traded in 2010. Saudi Arabia, China, and Japan were the
major Asian importers along with European countries Germany,
Belgium, and the Netherlands. Global domestic supply of barley
totaled to 140 Mt in 2009. Feed use accounted for 65% of barley
consumption while food use remained very low (4.5%).
Sorghum
The area under production for sorghum was 38 Mha in 2012
compared with 42 Mha in 2011 (Table 1). Global yield
increased by 151 kg ha 1 from 2011 to 1535 kg ha 1 in
2012. World sorghum production decreased 1 Mt from 2011
to 58 Mt in 2012. Nigeria, Mexico, the United States, India,
Argentina, China, and Ethiopia are the leading producers. Most
sorghum is produced by small-scale and subsistence farmers in
semitropical regions of Africa and Asia and by other farmers in
the United States and Latin America. Africa produced 40% of
the world total in 2012 on 23 Mha, more than half the world
total area. India has the largest area harvested for sorghum,
although there has been a significant reduction from 16 Mha in
1989 to 6.3 Mha in 2012. Production decreased in China from
a peak of 6 Mt in 1994 to 2 Mt in 2012 due to a decline in area
harvested. The United Arab Emirates had the largest average
yield in 2012 at 84 667 kg ha 1. World trade of sorghum was
13 Mt in 2010 with the United States and Argentina supplying
3.9 and 1.7 Mt to the export market, respectively. In 2010,
Mexico and Japan imported 2.3 and 1.6 Mt, respectively.
Mexico, the most important producer of sorghum at 7 Mt in
356 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview
2012, is also the largest importer. Chile, Ethiopia, the Republic
of the Sudan, and Spain are also main importers. Sorghum is
also an important commercial and export crop for Australia
and Argentina. The crop is utilized for feed (43%), food (44%),
and other uses (13%). Mexico, the United States, Australia,
Brazil, Japan, and Nigeria are principal feed users.
Millets
World production of all millets was 26 Mt in 2012 (Table 1),
46% of which was produced in Africa. West Africa (Nigeria,
Niger, Burkina Faso, Chad, Mali, and Senegal) produces 75%
of the millet output in Africa. Asia and Africa account for
96% of the global output of millet. India, China, Niger,
Burkina Faso, and Nigeria are the leading producers of millet.
Small-scale farmers produce almost all millet for household
consumption and localized trade. Very little millet (0.9 Mt or
3.5% of world millet production) is traded internationally.
India, the United States, China, Ukraine, and Austria are the
major exporters of millet. The European community accounts
for more than 23% of the global imports. Millet is primarily
dry-milled to produce cracked grain, grits, meal, and flour from
which a number of secondary and derived products are made.
Oats
World oat production in 2012 was 21 Mt, down by 2 Mt from
2011 due to a decrease in crop yields (Table 1). The total area
harvested was 10 Mha with an average yield of 2179 kg ha 1.
World oat production for 2012 by region, in Mt, was 7.9 (EU),
4.7 (FSU), 3.7 (NC America), 1.3 (Oceania), 0.6 (FE Asia), and
0.3 (ME Asia). Russia and Canada are the leading producers
followed by the United States and Australia. Canada is the
worlds largest exporter with the United States accounting for
over 90% of Canadas oat exports. The next largest export
regions are the EU (mainly Finland and Sweden) and Australia.
Russia generally consumes its feed quality oats domestically
even though it is the worlds largest single producer of oats. The
United States dominates global oat imports, accounting for
80%, followed by Germany, Mexico, and Japan. Oats is used
primarily for animal feed (71%) and human food (16%). High
levels of soluble fiber in oats have contributed to its promotion
as a healthy food ingredient.
Rye, Triticale, Buckwheat, Fonio, Quinoa,
and Canary Seed
World production of rye, triticale, buckwheat, fonio, quinoa,
and canary seed was 13, 14, 2.3, 0.6, 0.08, and 0.2 Mt,
Table 2 World pulse and oilseed production (Mt) for the period 201012
Year Total pulses Dry beans Dry peas Chickpeas
2010 70 23 10 11
2011 68 23 10 12
2012 70 23 10 11
Source: FAOSTAT Database 2013.
respectively, in 2012. These grains composed 1% of total
cereal production. Germany, Poland, the Russian Federation,
and Belarus are the major producers of rye. World buckwheat
production increased between 2010 and 2011 as area har-
vested and yields grew slightly. Decreases in area harvested
resulted in a decline in global canary seed production between
2010 and 2011. Worldwide production of both canary seed
and buckwheat remained steady between 2011 and 2012.
World quinoa production has steadily increased while global
triticale and fonio production has remained fairly constant
from 2010 to 2012. Canada contributed 62% of world
canary seed production in 2012. Canary seed is currently
used almost exclusively as birdseed. Rye, triticale, buckwheat,
fonio, and quinoa are used for human food mostly as ingredi-
ents in baked products. It is projected that the demand for
these grains will grow as whole grains continue to be promoted
as healthy foods.
Pulses
Table 2 gives data on world pulse production for the period
201012. Among pulses, beans were produced at 23.2 Mt
followed by chickpeas (10.9 Mt) and peas (10.2 Mt) in
2010. Asia contributes 45% of world pulse production.
Pulse production quadrupled in NC America and Africa since
1961. In the EU, pulse production peaked in 1990 at 7.4 Mt
and steadily declined to 3.2 Mt in 2012. India, Brazil, and
Myanmar are the leading producers of dry beans. The major
chickpea producer is India followed by Australia, Turkey,
Myanmar, and Ethiopia. Canada leads in both production
and export of dry peas in the world. Russia, France, China,
and India are also significant producers. World lentil produc-
tion is 4.6 Mt with Canada (1.9 Mt) and India (0.9 Mt) as
leading producers. India is a major importer of dry peas and
dry beans. Over 70% of pulse crops are consumed as human
food globally. Peas are used for both livestock feed and human
food. The proportion for the latter use is higher in Asia and
Latin America than in Europe.
Oilseeds
Major oilseeds produced include soybean, rapeseed, cotton-
seed, groundnuts, sunflower seed, palm kernels, and copra.
World oilseed production was 454 Mt in 2012 with soybean,
rapeseed/canola, cottonseed, groundnuts (unshelled), sun-
flower seed, and palm kernels making up 53, 13, 10, 8, 8.6,
and 3%, respectively, of the total production. World oilseed
production has risen tremendously since the 1970s due to the
Cow peas Pigeon peas Soybean Rapeseed
7 4 265 60
5 4 262 63
6 4 253 65
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview
expansion of the area planted. Soybean, palm, rapeseed, and
sunflower oil each accounted for 27, 32, 15, and 10,
respectively, of the world total oil (coconut, cottonseed,
groundnut, linseed, olive (virgin), palm, palm kernel, rape-
seed, safflower, sesame, soybean, and sunflower) produced
(155 Mt) in 2012. However, soybean made up about 70% of
the world protein meal consumption (187 Mt).
Soybean
World soybean production was 253 Mt in 2012 (Table 2)
compared with 26 Mt in 1961 (Figure 6). The United States,
Brazil, Argentina, China, and India accounted for 32, 26, 16,
5, and 4.5% of total production in 2012. In general, produc-
tion has been increasing since the 1950s due to the increase in
global area planted and crop yields; however, a decrease in
average yields between 2010 and 2012 caused a decline in
world soybean production. Soybean production is now world-
wide due to the crops ability to adapt to different types of soils
and climates and versatile end uses. In Europe, production
remains limited due to poor climate and soil conditions.
Argentina continues to expand soybean-processing capacity as
domestic policies continue to encourage value-added activities.
Its processing capacity was only half that of Brazil in 1999;
however, it has since surpassed that of Argentina with an
estimated 175 000 tonnes compared to 169 000 tonnes per
day in Brazil. Global soybean production is expected to
increase due to an increase in production in the Americas. A
growing demand for soybeans in China may lead to a growth
in world trade. World soybean exports were at 93 Mt in 2010.
Major exporters were the United States (45%), Brazil (28%),
Argentina (15%), Paraguay (4%), and Canada (3%).
Soybean is grown primarily for its meal in order to satisfy
the feed protein requirement of broilers, pork, and aquaculture
producers, which are not met by meat and fish meals and
meals from other oilseeds. Soybean oil is a secondary product.
Derived products (made from soybean, soybean meal, or soy-
bean by-product) include fermented foods (e.g., tofu, soymilk,
and soy sprouts) and nonfermented foods (soy sauce, miso,
tempeh, and natto). Soybean can be used to produce prod-
ucts differing in protein content. Other derived products
include soy ice cream, yogurt, burgers, cheese, meat analogs,
salad, cooking, and frying oils, and shortening and margarines.
300
250
Production (Mt)
200
150
100
50
0
1960 1970 1980
Figure 6 World soybean production for the period 19612012. FAOSTAT Database 2013.
357
Rapeseed/Canola
Rapeseed/canola production was 65 Mt in 2012 compared to
60 Mt in 2010 (Table 2). More than 88% of the worlds
rapeseed was produced in Asia, EU, and NC America, that is,
22, 19, and 16 Mt, respectively. China contributed 22% of total
rapeseed production. Canada produced 93% of the crop in NC
America. Canola is processed into vegetable oil for human
consumption and meal for livestock feed. Global rapeseed
production is expected to increase; however, this growth may
be limited by high prices and tight availabilities.
Exercises for Revision
Identify cereals classified as coarse grains and indicate
whether they are major or minor in terms of global
production.
Why are potential yields of wheat not yet realized in African
and Asian developing countries?
Which countries are the major exporters of wheat?
Globally, which regions or countries produce the most rice?
Which continent(s) identifies maize as the preferred crop?
What is the predominant use of barley?
Which countries are the leading producers of sorghum?
What are the major oilseed crops produced globally?
Exercises for Readers to Explore the Topic Further
Select one region and identify the major cereal crops pro-
duced. Are there variations in production among the lead-
ing producers over the last 5 years?
Oilseeds are important for the production of oil and meal
products. How does the current forecast differ from the past
year in production and consumption of soybean in North
America?
Maize is a preferred crop in most South American and
African countries. How is maize consumed in those
regions? Can sorghum replace maize to produce foods
consumed in those regions?
Are there minor cereal and oilseed crops that are likely to
become prominent in the near future in terms of produc-
tion and consumption? Give the rationale for your response
to this question.
1990 2000 2010
Year
358 GRAINS AROUND THE WORLD | Grain Production and Consumption: Overview
See also: Grains Around the World: Grain Production and
Consumption: Africa; Grain Production and Consumption: Cereal
Grains in North America; Grain Production and Consumption: Europe;
Grain Production and Consumption in Oceania: Australia and Pacific
Countries; Grain Production and Consumption: Oilseeds in North
America; Grain Production and Consumption: South America.
Further Reading
Agricultural Statistics (2000) National Agricultural Statistical Service. Washington:
US Government Printing Office; United States Department of Agriculture.
FAOSTAT (2013) FAO statistical databases. Food and Agriculture Organization of the
United Nations. Rome, Italy (website: http://faostat.fao.org).
Halweil B (2003) Grain production drops. In: Starke L (ed.) Vital Signs 2003. New York:
Norton.
IGC (2002) World Grain Statistics 2000/01. London: International Grains Council.
Signposts (2002) Envisioning the Future. Washington, DC: Worldwatch Institute.
A Worldwatch CD-ROM Resource.
Relevant Websites
http://www.angoc.ngo.ph Agrarian Reforms in Asia http://www.angoc.org/-Asian
NGO Coalition, for agrarian reform and rural development.
http://www.igc.org.uk International Grain Council market reports and grain statistics
Password required.
http://www.irri.org International Rice Research Institute world rice statistics.
http://www.soystats.com and http://www.unitedsoybean.org Oilseed and soybean
statistics.
http://www.fao.org and http://faostat.fao.org United Nations Food and Agriculture
Organization, probably the best single source of general articles, and FAOSTAT,
online databases covering international statistics in agriculture.
http://usda.mannlib.cornell.edu USDA Economics, Statistics and Market Information
System.
http://www.worldwatch.org Worldwatch research on food and agriculture.
 Grain Production and Consumption: Africa
JRN Taylor, University of Pretoria, Pretoria, South Africa
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Comprehensive information on the quantities of the vari-
ous grains (cereals, pulses, and oilseeds) produced in Africa
and where they are produced.
Review of Africas rich culture of traditional grain foods and
beverages.
Overview of the state of African agriculture and the causes
of low agricultural productivity.
Focus on the impact of rapid population growth and urban-
ization on food consumption and grain food security.
Information on the grain production and grain food supply
situation in Africa with respect to production and imports.
Review of major initiatives taking place in Africa to improve
agriculture and food security, including the application of
biotechnology and grain biofortification.
Learning Objectives
To know which are the important grains (cereals, pulses,
and oilseeds) produced in Africa and where they are
produced.
To understand the factors responsible for the generally low
agricultural productivity in Africa.
To be aware of Africas rich culture of grain foods and
beverages.
To understand the impact of rapid population growth and
urbanization in Africa on food security.
To be aware of the major initiatives taking place in Africa to
improve agriculture.
Introduction
Africa is the second largest continent with an area of 30.3
million square kilometers. It is the only continent that straddles
both tropics. Hence, Africa has a predominantly subtropical
and tropical climate. Most of North Africa and much of south-
western Africa have a desert climate, with < 250 mm of rainfall a
year. The climate in the African tropics varies from warm and dry
(rainfall 250500 mm year 1) to hot with a dry season or two
dry seasons (rainfall 5001000 mm) to the equatorial region,
which is hot with rain all year (10003000 mm). Only on the
northwestern Mediterranean seaboard, the southern Cape, and
South African eastern seaboard there is a mild, warm, and wet
climate (rainfall 5001000 mm). Africas natural vegetation
mainly ranges from desert and semidesert, though savanna, to
subtropical and tropical broadleaf forest. One significant excep-
tion is the central part of South Africa, which is natural grass-
land, the grassveld.
By 2015, Africas population would be 1.17 billion
(Figure 1). In the past 50 years from 1965 (the date when
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00043-7
most African countries had become independent), it has
grown exponentially, increasing 3.6-fold. The population is
still increasing rapidly but the rate of growth is slowing some-
what and is now  2.5% per year. Nevertheless, it is predicted
that by 2050, the population of sub-Saharan Africa will increase
to between 1.5 and 2 billion. Notably, half the worlds increase
in population to 2050 will be from sub-Saharan Africa.
As can be seen from Figure 1, both the rural and urban
populations are increasing, but urbanization is taking place
much more rapidly. It is predicted that by 2030, 50% of Africas
population will live in large towns and cities. Obviously, Africas
high population growth rate, coupled with rapid urbanization, is
currently having and will continue to have a profound impact on
the continents food security and food utilization.
This article examines the grain food situation in Africa, with
particular reference to the importance of grains as foodstuffs in
Africa, Africas major grains and their production, the causes of
grain agriculture low productivity, grain consumption, and the
impact of rapid population growth and urbanization and con-
cludes by reviewing developments and challenges in grain
agriculture and grain handling. The data are from the FAOSTAT
database of the Food and Agriculture Organization (FAO) of
the United Nations.
Importance of Grains
Africa is an exception compared to much of the rest of the
world in that quantitatively, cereals are only the second largest
foodstuff in terms of production. Starchy roots and tubers are
the largest, of which cassava accounts for just over half. The
quantities in 2012 were 172 million tonnes (Mt) of cereals
and  266 Mt of starchy roots and tubers. However, it should
be taken into account that cereal grains contain 70% starch,
whereas starchy roots and tubers only contain 35% starch,
and hence cereals are the larger contributor to energy intake.
Africa is also unusual in that only a relatively small proportion
of the cereals produced is used as animal feed, 12.5%. This
means that still today, the African diet is predominantly plant-
based. This is also reflected by the fact that the production of
plant protein, in the form of pulses,  15.9 Mt, is virtually
identical to that of meat, 16.4 Mt.
Grain Production
Cereal production in Africa is dominated by maize, which is a
tropical (C4) cereal, at 70 Mt (2012), some 40% of total
cereal production (Table 1). As can be seen, production is
very widespread across the continent both geographically and
climatically. In fact, maize production is recorded for 50 coun-
tries in Africa. However, the top four producing countries
South Africa, Nigeria, Egypt, and Ethiopia are responsible for
about half of the total production.
359
360 GRAINS AROUND THE WORLD | Grain Production and Consumption: Africa

1200

1000

Population (millions)
800 Total

600
Rural

400
Urban

200

0
1965 1970 1975 1980 1985 1990 1995 2000 2005 2010 2015
Year
Figure 1 Africas population growth from 1965 to 2015. Data from FAOSTAT: http://faostat.fao.org. Population (accessed 21.04.14.).

Table 1 Grains produced in Africa (2012 data)

Production Area harvested Yield Major producing countries in descending

Grain (t  103) (h  103) (t ha 1) order

Cereals
Cereals (total) 172 184 106 955 1.61 Nigeria, Egypt, Ethiopia, South Africa
Barley 6023 (3.5)a 4997 1.21 Ethiopia, Algeria, Morocco
Fonio 587 (0.3) 566 1.04 Guinea, Nigeria
Maize 69 637 (40.4) 33 710 2.08 South Africa, Nigeria, Egypt, Ethiopia
Millet (mainly pearl millet) 16 083 (9.3) 19 978 0.81 Nigeria, Niger, Mali
Oats 276 (0.2) 199 1.39 Algeria, South Africa, Morocco
Rice (paddy) 26 824 (15.6) 10 538 2.54 Egypt, Nigeria, and Madagascar
Rye 94 (<0.1) 52 1.80 Egypt
Sorghum 23 313 (13.5) 23 169 1.01 Nigeria, Ethiopia, Burkina Faso, and Sudan
(former)
Triticale 26 (<0.1) 13 2.01 Tunisia
Wheat 24 711 (14.4) 10 217 2.42 Egypt, Morocco, Ethiopia, and Algeria
Pseudocereals
Buckwheat 10 11 0.97 Tanzania
Pulses (grain legumes)
Pulses (total) 15 865 24 447 0.65 Ethiopia, Nigeria, Tanzania, and Niger
Bambara bean 159 (1.0)b 233 0.68 Burkina Faso, Niger, Cameroon, and
Mauritania
Dry beans (common beans) 5070 (32.0) 7829 0.65 Tanzania, Kenya, Ethiopia, and Rwanda
Broad bean 1505 (9.5) 670 1.55 Ethiopia
Chickpea 697 (4.4) 595 1.17 Ethiopia
Cowpea 5403 (34.1) 11 043 0.49 Nigeria, Niger
Lentil 187 (1.2) 188 0.99 Ethiopia
Peas (dry) 596 (3.8) 715 0.83 Ethiopia and Tanzania
Pigeon pea 632 (4.0) 754 0.84 Malawi and Tanzania
Pulses (other) 1262 (8.0) 1890 0.67 Mozambique, Sudan (former)
Oil-rich legumes
Peanuts (groundnuts) 10 942 11 823 0.93 Nigeria, Sudan (former), and Tanzania
(in shell)
Soybean 1986 1638 1.21 South Africa and Nigeria
Oilseeds
Rapeseed 201 113 1.79 South Africa, Ethiopia, and Algeria
Sunflower seed 2224 1780 1.24 Tanzania and South Africa
a
Percentage of cereal production.
b
Percentage of pulse production.
Data from FAOSTAT http://faostat.fao.org. Production (accessed 21.04.14.).
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Africa
Quantitatively, the next most important cereals, in descend-
ing order, are rice, wheat, and sorghum, accounting for 15.6%,
14.4%, and 13.5% of total cereal production. Production of
these three cereals is also widespread across Africa, with 43
producing countries being recorded for rice, 41 for sorghum,
and 31 for wheat. However, there are some important differ-
ences with respect to their cultivation. Africas production of
sorghum, 23 Mt, is  40% of world sorghum production.
The area where sorghum is harvested in Africa, 23 M ha, is 60%
of the world total. The major production is in the semiarid
tropics of North Africa, from Nigeria and Burkina Faso in the
west to Sudan and Ethiopia in the east (Table 1). This is
probably on account of the fact that sorghum, a tropical (C4)
cereal, is indigenous to Africa and is very well adapted to harsh
climatic conditions, being able to withstand periods of drought
and also waterlogging. Sorghum requires a minimum of only
400 mm of water for cultivation, as opposed to the
500600 mm needed by maize. In contrast, the production
of rice (a tropical C3 cereal) requires much more water and is
concentrated in areas of high water availability: the Nile valley
of Egypt, tropical Nigeria, and Madagascar. Concerning wheat,
which is a temperate C3 cereal, its production is generally
highest in countries in Africa on the Mediterranean coast. The
top three producers are Egypt, Morocco, and Algeria. Substan-
tial quantities of wheat are also grown in countries in the
tropics, in particular Ethiopia and Kenya, with cultivation
being carried out at elevated altitudes to attain the required
cooler conditions.
Africa is the home to a number of millet species, which are
all C4 cereals. Millets are, by definition, small grained cereals.
The FAO data (Table 1) only specifically record the millet
species fonio (Digitaria sp.), also known as acha, or more
colloquially as hungry rice on account of its ability to produce
a crop under extreme hot and dry conditions. Fonio is endemic
to the Sahel region of West Africa, in particular Guinea
(Table 1). The other millets are lumped together under the
general heading millet. Millet production in total accounts for
19% of the land area under cereal cultivation in Africa. By far,
the most important millet is pearl millet (Pennisetum glaucum),
accounting for  87% of millet production in Africa and  8%
of the continents total cereal production. Pearl millet is
uniquely able to produce a crop under very low rainfall condi-
tions, with a minimum water requirement of only 300 mm
and very high temperatures (daytime temperatures above
30  C). Pearl millet production is also mainly in the dry areas
of West Africa, but it is also cultivated widely across the conti-
nent. The other economically important millets in Africa in
descending order of production are finger millet (Eleusine cor-
acana), teff (tef) (Eragrostis tef), and guinea millet (Brachiaria
deflexa). Finger millet, so-called because the grains are borne
on fingerlike panicles (Figure 2(a)), is grown throughout sub-
Saharan Africa. It is an especially important grain with regard
to rural food security on account of its excellent storability,
being resistant to insect attack on account of its proportionally
small germ. Teff production is almost exclusively in Ethiopia
and Eritrea.
There is also substantial production of barley in Africa,
which is produced both for brewing and as a food, the latter
especially in Ethiopia. Production of oats, rye and triticale, and
the pseudocereal buckwheat is very limited (Table 1).
361
Figure 2 Finger millet and bambara beans. (a) Finger millet under
cultivation in Kenya, (b) Bambara beans from Namibia, (c) Bambara
beans from Zimbabwe (close up).
As mentioned, pulses are a major food crop in Africa. Total
pulse production is 16 Mt per year and is dominated by
cowpea (Vigna unguiculata) and dry beans (Phaseolus vulgaris),
the production of both being 5 M tonnes per annum (Table 1),
and together, they account for  66% of pulse production.
Cowpea is indigenous to Africa. The major producing countries
are in tropical West Africa, in particular Nigeria and Niger,
although production is widespread throughout Africa with sig-
nificant production recorded in 19 countries. Dry beans (also
known as common beans) originated in South America. The
major producing countries are in East Africa, with the highest
production in Tanzania, Kenya, and Ethiopia. However, there is
significant production across the continent in at least 33 coun-
tries. Broad bean (Vicia faba) is quantitatively the third most
important bean with an annual production of 1.5 M tonnes,
with Ethiopia responsible for 63% of Africas production. The
production of other pulses, including chickpea (Cicer arietinum),
pigeon pea (Cajanus cajan), dry pea (Pisum sativum), lentil (Lens
culinaris), bambara bean (Vigna subterranea), and other exotic
and indigenous pulses such as lablab (Lablab purpureus), yam
bean (Pachyrhizus sp.), mung bean (Vigna radiata), and marama
bean (Tylosema esculenta) is also of significant importance espe-
cially with regard to rural food security and in the case of dry peas
and lentils as cash crops.
Of particular interest is the bambara bean (Figure 2(b) and
2(c)), also known as the African or bambara groundnut, which
is indigenous to Africa. It is widely cultivated across the drier
parts of topical Africa, with, according to one authoritative
source, Zimbabwe being the center of production, whereas
FAOSTAT only records significant production in five countries,
all in the Sahel region, with Burkina Faso being the largest
producer. The bambara bean can be cultivated where temper-
atures reach as high as 40  C and is one of the most drought-
tolerant legume species. It is considered to be one of the
worlds most underestimated and neglected crop plants.
Total production of oil-containing grains (oilseeds) in Africa
is also  16 Mt per year and is dominated by peanuts (ground-
nuts) at 10.9 Mt (Table 1). This is related to the fact that
peanuts are used as both a food and a source of cooking oil.
362 GRAINS AROUND THE WORLD | Grain Production and Consumption: Africa

Production takes place throughout Africa, with 47 countries
recorded as significant producers. However, Nigeria produces
by far the largest quantity of peanuts,  3.1 Mt. Significant
quantities of oil-bearing grains other than peanuts are also
produced in Africa. The most important are sunflower and
soybean, with a total production of  2.2 and 2.0 Mt, respec-
tively. There is also some production of rapeseed (canola).
South Africa is the major producer of both soya and canola.
Grain Agriculture
Figure 3 shows the trends in cereal production and imports for
Africa over a 45-year period since the mid-1960s when most
African countries gained independence, to 2010. Over the
period, the area harvested remained more or less constant up
until the about 1980 and then steadily but gradually increased,
with an  73% increase over the entire period. In contrast,
yield initially increased over the first 10 years and then
remained static until the mid-1990s and has subsequently
increased substantially in recent years. Over the whole period,
yields increased by  83%. Increased area of harvest and
increased yields have resulted in a continuous increase in cereal
production, which has accelerated since the mid-1990s. Over-
all, cereal production increased 3.2-fold, keeping pace with
but not exceeding the increase in population 3.1-fold. As a
consequence, Africas cereal imports have grown at a rapid but
linear rate over the period, increasing from 11% to 34% of
cereal production. Imports of maize now represent some 26%
of local production (Table 2).
The need to import an increasing proportion of cereal
grains is related to the fact that much of Africas agriculture is
still subsistence farming. However, small-scale commercial and
large-scale mechanized farming are widespread and increasing.

180 1.80

160 1.60
Production/area harvested/imports (Mt)

140 1.40

120 1.20

Yield (tonnes/Ha)
100 Yield 1.00

80 Area 0.80
Production harvested
60 0.60

40 0.40
Imports

20 0.20

0 0.00
1965 1970 1975 1980 1985 1990 1995 2000 2005 2010
Year
Figure 3 Africas cereal production and imports from 1965 to 2010 (data from FAOSTAT5). Data from FAOSTAT. Production: Crops http://faostat.fao.
org/ (accessed 21.04.14.).

Table 2 Africas domestic grain-based food and beverage supply (2009) (t  103)

Imports (% of Exports (% of Stock Domestic Domestic supply

Products Production Imports production) Exports production) changes supply (% of production)

Cereals (total) 146 217 55 987 38.3 4065 2.8 1163 199 303 136.3
Maize 57 702 14 995 26.0 2132 3.7 3498 67 067 116.2
Wheat 26 074 30 288 116.2 897 3.4 126 55 339 212.2
Barley 5827 1744 29.9 29 0.5 306 7237 124.2
Lager and stout beers 9758 510 5.2 188 1.9 199 9882 101.3
Pulses (total) 12 086 1262 10.4 577 4.8 322 12 803 105.9
Peanuts (groundnuts) 9604 227 2.4 148 1.5 16 9670 100.7
(in shell)
Soybeans 1612 1133 70.3 192 11.9 170 2722 168.9
Sunflower seed 1689 175 10.4 14 0.8 116 1734 102.7
Groundnut oil 1374 9 0.7 48 3.5 16 1319 96.0
Soybean oil 300 1449 483.0 127 42.3 87 1535 511.7
Sunflower seed oil 597 596 99.8 90 15.1 132 971 162.6

Data from FAOSTAT http://faostat.fao.org. Production (accessed 21.04.14.).

 GRAINS AROUND THE WORLD | Grain Production and Consumption: Africa
Figure 4 Grain harvesting in Africa. (a) Manual harvesting of teff in
Ethiopia. (b) Combine harvesting of wheat in Kenya.
Unfortunately, the so-called green revolution has barely
impacted on the subsistence farming sector. Here, agriculture
is characterized by traditional farming practices using manual
labor and some draft animal power (Figure 4(a)), low inputs
(no inorganic fertilizer or pesticides), and the use of low-
yielding traditional varieties or landraces. Taking the example
of sorghum, which in Africa is mainly produced by subsistence
farmers, yields have only increased marginally over the period
from 0.75 to just 1.0 t ha 1. This compares to the sorghum
yield of almost 4 t ha 1 in the United States where mechanized,
high-input agriculture using hybrid seeds is employed. Where
these technologies are also applied in Africa, for example, in
South Africa, yields are substantially higher,  2.7 t ha 1.
Also, as can been seen from Table 1, agricultural produc-
tion efficiency in Africa varies very considerably between the
grains, with a low of millet at 0.8 t ha 1 among the cereals, and
an overall low of only 0.5 t ha 1 for cowpea, and a high of
2.4 t ha 1 for wheat among the cereals. Production efficiency
appears to depend, to a considerable extent, on whether the
grain is primarily home-processed as in the case of cowpeas or
industrially processed as in the case of wheat. In other words,
production is driven by the existence and needs of a linked
food processing industry, which encourages the development
of larger-scale mechanized farming, as, for example, has taken
place with wheat in Kenya (Figure 4(b)). Hence, the average
yield of wheat in Africa of 2.4 t ha 1 compares reasonably well
with the  3.1 t ha 1 in the United States of America.
Grain Food Consumption
Africa has a rich culture of traditional grain foods and bever-
ages (Figure 5). The following gives some idea of the huge
variety of cereal and legume products:
Roasted snack foods, for example, kollo barley
(Ethiopia)
Leavened baked wheat flour breads, for example, geish
baladi (Egypt)
363
Leavened steamed wheat flour breads, for example,
ledombolo (South Africa)
Flatbreads, for example, kisra sorghum or millet (Sudan)
Pancakes, for example, injera tef, sorghum, finger millet,
wheat or maize, or combinations (Ethiopia)
Dumplings, for example, kenkey maize (Ghana)
Whole or dehulled boiled grains, for example, supa
mtama sorghum (Kenya)
Steamed, granulated foods, for example, couscous
wheat, sorghum, or pearl millet (North and West Africa)
Stiff porridges, for example, sadza, maize (Zimbabwe),
and ugali, maize, sorghum, and finger millet (East
Africa)
Soft porridges and gruels, for example, uji maize, sor-
ghum, or finger millet (East Africa)
Porridge cooked with wood ash extract making it alkaline,
for example, to sorghum or pearl millet (Mali)
Porridge, lactic acid-fermented, making it sour, for exam-
ple, ogi, maize, sorghum, or pearl millet (Nigeria), and
ting, sorghum (Botswana and South Africa)
Porridge flavored with tamarind or lemon juice, making it
acidic, for example, to sorghum (Burkina Faso)
Traditional African cereal beverages are both nonalcoholic, for
example, mageu (southern Africa), and alcoholic, for exam-
ple, cloudy pito beer (Ghana) and opaque sorghum beer
(central and southern Africa). The beverages are generally opa-
que and viscous in consistency, due to the presence of semi-
suspended starch. They are produced variously from sorghum,
maize, pearl millet, and finger millet, either singly or in com-
bination. In the beers, a portion of the cereal is in the form of
malt, in order to provide amylase enzymes to hydrolyze the
starch into fermentable sugars.
A characteristic of many traditional African cereal foods and
beverages is that they have undergone a lactic acid bacterial
fermentation during processing. This gives the product a char-
acteristic sharp, sour taste and helps preserve it against micro-
bial spoilage. Examples include the pancake injera, the
dumpling kenkey, the firm porridge ting, the thin porridge
ogi, the nonalcoholic beverage mageu, and sorghum beer.
Legume-based traditional African foods include
blanched and roasted whole grain snacks, for example,
kollo, chickpea (Ethiopia);
boiled whole grain snacks, for example, nifro, chickpea,
broad bean, or lentil (Ethiopia);
boiled whole grains foods, for example, dikgobe, cowpea
(Botswana); mayengele, dry beans (Uganda); and
mtakura, cowpea, peanuts, or bambara bean (Zimbabwe),
all often mixed and served with boiled maize grains;
steamed pastes, for example, moi-moi, dry beans
(Nigeria), and okpa, bambara bean (Nigeria);
deep-fried pastes, for example, kose, cowpea or dry beans
(Ghana);
deep-fried bean balls, for example, akara, dry beans
(Nigeria);
fermented pastes, for example, siljo, broad bean
(Ethiopia);
boiled grains made into a sauce, for example, magila,
cowpea (Uganda);
germinated and boiled grains made into a sauce, for exam-
ple, azifa, chickpea, broad bean, or lentil (Ethiopia); and
364 GRAINS AROUND THE WORLD | Grain Production and Consumption: Africa

Figure 5 Selection of African grain foods and beverages. (a) Kisa flatbread (sorghum) Sudan, (b) to porridge (maize) with chickpea relish and
goat meat Burkina Faso, (c) Ugali porridge (sorghum) Kenya, (d) Baladi bread (wheat) Sudan, (e) Fonio couscous and peanut sauce (Burkina Faso),
(f) Mealie bread (maize and wheat) South Africa, (g) Samp and beans (maize and dry beans) southern Africa, (h) Opaque sorghum beer
(maize and sorghum) southern Africa, (i) Mageu fermented nonalcoholic beverage (sorghum and maize) Zimbabwe and South Africa.

dry roasted grain milled into powder, spiced, and made
into a sauce, for example, shiro, pea, broad bean, or chick-
pea (Ethiopia).
However, food consumption patterns in Africa are changing
rapidly as a result of urbanization. The so-called nutrition
transition is resulting in a dramatic increase in consumption
of Western-style convenience foods, including bread, potato
chips (French fries), fried chicken, and beef burgers, and bev-
erages, in particular carbonated soft drinks and lager beer. For
example, with regard to convenience foods, the largest South
African fast-food franchise company opened 18 restaurants in
other African countries in the last quarter of 2012. In the case
of lager beer, the figures are if anything even more dramatic.
From 2007 to 2012, the largest brewing company in Africa
recorded an average annual 12% increase in beer volume.
These changing food consumption patterns are severely
impacting on the grain food supply situation in Africa. With
regard to cereals, the continent is importing as much wheat as
is produced locally (Table 2). In the case of barley, imports
amount to 30% local production. The situation with regard
to soybeans, the most important protein source in formulated
animal feeds, is worse, with domestic supply being  170% of
production. With regard to seed oils, which are majorly used to
manufacture plant-based fats like margarine, the situation is
even more dire. Africa is importing nearly five times the quan-
tity of soy oil it produces and as much sunflower seed oil as it
produces. Taking the previously mentioned into account, the
fact that Africa is essentially self-sufficient in lager and beer
production may seem surprising. Part of the explanation is that
a substantial proportion of these beers are now brewed using
locally grown sorghum as either the sole or major ingredient,
especially in West and East Africa.
Developments and Challenges in Grain Agriculture
and Grain Handling
The high dependence on imported grains has a very adverse
effect on the fragile economies of many African countries. To
prevent increasing poverty and malnutrition resulting from
rapid population growth and urbanization, it is essential that
the effectiveness of African grain agriculture is improved. How-
ever, the improvement of agriculture in Africa faces a number
of severe challenges. These include the acute need to introduce
modern agricultural production technologies and chronic
environmental problems such as decreasing soil fertility and
desertification, resulting from farming more marginal land.
There are also sociopolitical issues. Governmental policies are
often not conducive to local agricultural development. As of
2013, only eight countries in Africa had met the UN Millen-
nium Development Goal of halving the proportion of hungry
people. Persistent problems include keeping food prices artifi-
cially low to subsidize the urban consumer, but at the cost of
the farmers livelihood, and poorly planned and executed land
reforms that disrupt large-scale commercial farming
operations.
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Africa 365

The application of biotechnology to grain agriculture in

Africa has the potential to provide many benefits that could
help the continent leapfrog the first green revolution. Geneti-
cally modified (GM) crops are currently being commercially
cultivated in some countries, notably South Africa and Burkina
Faso. For example, some 80% of maize produced in South
Africa is GM glyphosate herbicide-tolerant and/or Bt gene
insect-resistant. It is stated that latter technology has increased
yields by 11%. However, some African countries refuse to even
import GM grains, never mind grow them. Notwithstanding
this, GM plants with built-in specific resistance to pests,
tolerance to environmental stresses such as drought and
low rainfall, and improved nutritional value to combat
malnutrition should be of tremendous value.
Biotechnology alone will not be a panacea. In recent years,
there have been several major collaborative initiatives to
improve Africas agriculture, notably the Alliance for a Green
Revolution in Africa (AGRA). AGRA aims to develop practical
solutions to significantly boost farm productivity and incomes
for the poor while safeguarding the environment. Perhaps the
most important of AGRAs projects is the Program for Africas
Seed Systems (PASS). PASS supports in-country-breeding
teams who work closely with farmers to develop new varieties
and with local entrepreneurs to establish independent seed
companies to produce and distribute the seed. AGRA also
supports the training of high-level agricultural scientists, for
example, training PhD-level plant breeders through the African
Centre for Crop Improvement, hosted by the University of
KwaZulu-Natal in South Africa.
A notable initiative is the development of biofortified staple
crops for smallholder farmer cultivation in Africa. Biofortifica-
tion is the enhancement of the levels of critically deficient Figure 6 Grain handling challenges in Africa. (a) Outdoor storage of
nutrients, particularly micronutrients, in staple food crops sorghum at harvest time in Burkina Faso. (b) Poorly cleaned smallholder
such as grains. It can be achieved by conventional breeding farmer-produced sorghum being unloaded at a factory in Nigeria.
and also by recombinant DNA technology (genetic engineer-
ing). Probably the most well-known biofortified crop is quality
protein maize (QPM), the protein of which is much richer in produced grain is often more expensive than imported grain.
lysine and other indispensible amino acids than conventional Additionally, there can also be severe problems with quality
maize. Several studies have shown that consumption of QPM that render small-scale farmer-produced grain unsuitable for
maize has improved the nutritional status of young children, industrial processing. Consignments of grain may comprise
for example, in Ethiopia. Recent biofortified cereals include different varieties with differing grain properties such as grain
provitamin A maize, iron- and zinc-rich pearl millet, and Africa size, and worse, the grain may be heavily contaminated with
Biofortified Sorghum. The latter, which has been developed by dirt and stones (Figure 6(b)).
genetic engineering, is unique in that the final product will
have enhanced multiple nutrients: provitamin A, improved
protein quality and protein digestibility, and increased iron
and zinc bioavailability. Exercises for Revision
As Africa urbanizes and develops its grain processing indus-
tries, a further challenge that still needs to be properly State in descending order which are quantitatively the
addressed is for countries to dramatically improve their grain major cereals, pulses, and oilseeds produced in Africa.
handling systems. Lack of adequate grain storage facilities in Give examples of indigenous African cereals and pulse
many countries causes ruinously low prices for farmers when grains (at least three of each) and state where they are
crops are good and acute shortages for consumers when crops cultivated.
are poor (Figure 6(a)). The almost complete absence of rail- Give examples of traditional African grain foods and bever-
ways and lack of good roads in much of Africa mean that the ages (at least three of each) and state what type of product
costs of transporting grain from the point of production to they are and from which grain(s) they are made.
where it is processed are very high. This is compounded by State which are the major grains and grain products that
high costs of grain assembly, where many small consignments Africa imports.
of grain from different small-scale farmers have to be brought Explain the reasons for the generally poor agricultural pro-
to one place to make up a truckload. As a result, locally ductivity in Africa.
366 GRAINS AROUND THE WORLD | Grain Production and Consumption: Africa
Explain what is the impact of rapid population growth and
urbanization on food security in Africa.
Explain the concept of biofortification and its benefits.
Exercises for Readers to Explore the Topic Further
Improving smallholder agriculture productivity in Africa
using appropriate technologies.
Improving grain storage using appropriate technologies.
Reduction in the cost of assembly and transport of consign-
ments of grain produced by smallholder farmers.
Development of simple but effective grain-quality stan-
dards suitable for smallholder farmers.
Development of technologies to produce inexpensive but
good-quality bread using maize and sorghum.
Further Reading
Akalu G, Taffesse S, Gunaratna NS, and De Groote H (2010) The effectiveness of quality
protein maize in improving the nutritional status of young children in the Ethiopian
highlands. Food and Nutrition Bulletin 31: 418430.
National Research Council (1996) Lost Crops of Africa. vol. 1: Grains. Washington, DC:
National Academies Press.
National Research Council (2006) Lost Crops of Africa. vol. 2: Vegetables. Washington,
DC: National Academies Press.
Popkin BM (2003) The nutrition transition in the developing world. Development Policy
Review 21: 581597.
Rohrbach DD (2003) Improving the commercial viability of sorghum and pearl millet in
Africa. Afripro Paper 22. http://www.afripro.org.uk (accessed 21.04.14).
Saltzman A, Birol E, Bouis HE, et al. (2013) Biofortification: Progress toward a more
nourishing future. Global Food Security 2: 917.
Taylor JRN, Dlamini BC, and Kruger J (2013) 125th Anniversary Review: The science of
the tropical cereals sorghum, maize and rice in relation to lager beer brewing.
Journal of the Institute of Brewing 119: 114.
Van Wyk B-E and Gericke N (2000) Peoples Plants: A Guide to Useful Plants of
Southern Africa. Pretoria: Briza.
Relevant Websites
http://agra-alliance.org AGRA (accessed 21.04.14.).
http://www.fao.org FAO. 12 June 2013. 38 countries meet anti-hunger targets for
2015 (accessed 21.04.14.).
http://faostat.fao.org/ FAOSTAT. Commodity Balances: Crops Primary Equivalent
(accessed 21.04.14.).
http://faostat.fao.org FAOSTAT. Population (accessed 21.04.14.).
http://faostat.fao.org/ FAOSTAT. Production: Crops (accessed 21.04.14.).
http://www.financialmail.co.za Financial Mail. 6 July 2012. SABMiller barrels ahead
(accessed 21.04.14.).
http://www.fao.org How to Feed the World 2050: High-level Expert Forum. The
special challenge for sub-Saharan Africa (accessed 21.04.14.).
http://www.howwemadeitinafrica.co.za How We Made It In Africa. More than enough
potential for restaurants and fast-food in Africa (accessed 21.04.14.).
http://education.nationalgeographic.com National Geographic Education. Africa:
Physical Geography (accessed 21.04.14.).
http://www.pannar.com PANNAR Seed. 2013 Product Catalogue (accessed 21.04.14.).
http://unhabitat.org UN Habitat: The State of African Cities, 2012 (accessed 21.04.14.).
 Production and Consumption of Grains: India
BS Khatkar, N Chaudhary, and P Dangi, G. J. University of Science and Technology, Hisar, India
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Agriculture accounts for considerable importance in Indias
economic development, as it provides food for more than
1.2 billion people and total employment to about 54.6% of
the population.
India holds the second-largest agricultural land (179.9
million hectares) in the world.
Food grain production covers the dominant part of the
cropped area (65%) in Indian agriculture.
India is the worlds largest producer of millets and second-
largest producer of wheat, rice, and pulses.
The highest food grain production of 259.32 million
tonnes was recorded in 201112.
The cereals other than rice and wheat contributed 1 kg per
person per month in rural India and 0.8 kg per person per
month in urban India in 201112.
The consumption of all food grains per person in the rural
areas is projected to decline from 15.3 kg per month in
2000 to 13.8 kg per month by 2050 and only slightly
decrease from 11.8 to 11.6 kg per month in the urban areas.
The nongrain food crops and animal products contributed
27% and 8%, respectively, of the total caloric requirement.
Meat consumption in India is very low due to cultural and
religious reasons, and most of the calories from animal
products are obtained from milk and milk products.
Learning Objectives
To present past and future prospects of the Indian food
grain production.
To identify the role of the Indian government in supporting
agriculture and food grain production.
To understand food grain consumption patterns of the
Indian population.
To learn the status of changing food grain consumption
patterns and the reasons behind these shifts.
Introduction
India accounts for only about 2.4% of the worlds geographical
area and 4% of its water resources, but has to support about
17% of the worlds human population and 15% of the live-
stock. India holds the second-largest agricultural land (179.9
million hectares) in the world. Food grain production covers
dominant part of the cropped area (65%) in Indian agriculture.
According to the FAO world agriculture statistics (2010), India
is the worlds largest producer of millets and second-largest
producer of wheat, rice, and pulses.
Indian agriculture has witnessed wide ups and downs in
growth performance during the last six decades. Performance
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00044-9
of Indian agriculture can be divided into six phases viz. pre-
green revolution period (196061 to 196869), early green
revolution period (196869 to 197576), period of wider
technology dissemination (197576 to 198889), period of
diversification (198889 to 199596); postreform period
(199596 to 200405), and period of recovery (200405 to
201011). The green revolution period was the golden period
for Indian agriculture, as it witnessed tremendous growth in
agricultural production. During the period of wider dissemi-
nation of technology, the spread of green revolution technol-
ogies pushed to maintain the increasing rhythm of growth
across various regions realized during the previous period.
However, a visible deceleration of growth was experienced
during the postreform period, as both public and private
investments suffered a setback, which resulted in the sluggish
performance of the sector. Moreover, yield levels of most of the
crops suffered a deceleration because use of primary resources
in the sector also slowed down. The descending trend of
growth continued up to 200405, after which a sharp return
was attained that can be attributed to a conscious increment in
public and private ventures and substantial improvement in
trade in favor of the agricultural sector.
Food Grain Production
India is continuously facing pressure on the demand side due
to steady population growth, limited land availability, and
several other production deterrents, which might also appear
as obstacles for the supply of food grains. Indian government
policies and planning has always given considerable impor-
tance to production of food grains due to which India has been
achieving the continued growth despite many constraints. Dur-
ing the last decade (200001 to 201011), the areas under
cereals, pulses, and oilseeds have increased by about 5.7, 6.0,
and 4.0 million hectares, respectively, and the area under
coarse cereals has declined by 2.6 million hectares. The area
under wheat cultivation has accelerated significantly in com-
parison to other cereals and millets. However, a moderate
deceleration in total grain cultivated area in comparison to
pulses cultivated area was witnessed during the years of 2005
to 2011. The highest food grain production of 259.32 million
tonnes was recorded in 201112. Out of the total food grains
production, cereals accounted for 242.23 million tonnes and
pulses 17.09 million tonnes (Figure 1). Production of rice and
wheat was recorded at 105.3 and 94.9 million tonnes, respec-
tively. It was estimated that the agriculture sector in India is
likely to grow by 5.25.7% in 201314. India is one of the
largest producers of cereal as well as the largest exporter of
cereal products worldwide. Indias export of cereals during
201314 stood at Rs. 63452.09 crores. Rice (including basmati
and non-basmati) occupies the most sizable share in Indias
total cereals export with 64.40% during this period. Countries
367
368 GRAINS AROUND THE WORLD | Production and Consumption of Grains: India

Total Pulses Coarse Cereals Total cereals Total Foodgrains

300

Production (million tonnes)

250

200

150

100

50

0
2010-11 2011-12 2012-13
Year
Figure 1 Production of food grains in India.

40
33.8
35
Area (percentage)

30
25
20
14.8
15 12.7
9.7 8.7
10 7.8
3.5 2.8
5 1.8 1.5 1.5 1.4
0
ra

at

sh
sh

ab

na

s
ha

ga
ha

er
es

ha

ar
ht

de
de

ya
nj

nc

en

th
Bi
ad

st

uj
as
Pu

ar

ra
ra

O
ra

tB
ja

G
Pr

ar
H

lP
rP

Ra

tta
ah

es
a

ha
tta

U
hy

ac
U

ad

im
M

States
Figure 2 Share of states in India in terms of area under cultivation of wheat.

such as Iran, Saudi Arabia, Indonesia, UAE, and Bangladesh Uttaranchal, Himachal Pradesh, and Jammu and Kashmir
mainly imported cereals from India during 201314. (Figure 3). These states contribute about 99.5% of total
wheat production in the country. The remaining states, namely
Jharkhand, Assam, Chhattisgarh, Delhi, and other
Wheat northeastern states, contribute only about 0.5% of the total
The area and production under wheat has increased from wheat production in the country. Among food grains, wheat in
27.99 million hectares in 200607 to 29.86 million hectares India stands next to rice both in area and production.
in 201112 and from 75.81 million tonnes in 200607 to an
all-time record high of 94.88 million tonnes in 201112,
Rice
respectively. Figure 2 shows the percent share of states in
India in terms of area. Uttar Pradesh, Madhya Pradesh, Punjab, India is the second-largest producer and consumer of rice in
Rajasthan, and Haryana are the leading states in terms of area the world and accounts for 22.3% of global production. Sev-
under wheat cultivation. Uttar Pradesh accounts for the maxi- eral programs run by the government such as National Food
mum area under cultivation and production, whereas Punjab Security Mission (NFSM) launched during 200708 and Bring-
and Haryana are yielding higher productivity with lesser land- ing Green Revolution in Eastern India (BGREI) during
holding under wheat cultivation. 201011 increased production and productivity of rice from
The increase in production of wheat has been observed in 96.7 million tonnes and 2202 kg ha 1 in 200708 to 105.31
the states of Haryana, Punjab, Madhya Pradesh, and Uttar million tonnes and 2393 kg ha 1, respectively, in 201112.
Pradesh. The normal national productivity is about 2.9 The NFSM is presently under implementation in 482 dis-
tonnes per hectare. The major wheat-producing states are tricts of 19 states of the country with a view to enhance the
Uttar Pradesh, Punjab, Haryana, Madhya Pradesh, Rajasthan, production of rice, wheat, and pulses through area expansion
Bihar, Maharashtra, Gujarat, Karnataka, West Bengal, and productivity enhancement, restoring soil fertility and
 GRAINS AROUND THE WORLD | Production and Consumption of Grains: India 369

40
33.7
35

Production (percentage)
30
25 21.2
20
15 12.9
9.6 8.9
10 6.2
5 1.7 1.6 1.2 1 0.8 1.2
0

ab

na

ra

al

sh

rs
at

ga
ha
es

es

ha

ch

e
ht

ar

de
ya
nj

en

th
Bi
ad

ad

st

as

uj

an
Pu

ar

ra

O
tB
ja

G
Pr

Pr

ar

r
H

lP
Ra

tta
ah

es
r

ha
tta

U
hy

ac
U

ad

im
M

H
States
Figure 3 Share of states in India for production of wheat.

25
Production (million tonnes)

20

15

10
5

4
-0

-0

-0

-0

-0

-1

-1

-1

-1

-1
04

05

06

07

08

09

10

11

12

13
20

20

20

20

20

20

20

20

20

20

Year
Figure 4 Status of maize production in India.

productivity, creating employment opportunities, and enhanc- improving, and its cultivation has spread to other parts of
ing farm level economy to restore confidence of farmers. The Uttar Pradesh and Madhya Pradesh.
basic strategy of the mission is to promote and extend
improved technologies that is, seed, micronutrients, soil
Maize
amendments, integrated pest management, farm machinery,
and resource conservation technologies, along with capacity After rice and wheat, maize is emerging as the third-most-
building of farmers with effective monitoring and better man- important crop in India. Its importance lies in the fact that it
agement in the high-potential districts to bridge the yield gaps. is used for human food as well as animal feed, and it is also
BGREI aims to increase rice productivity by intensive culti- widely used for the starch industry, oil production, baby corns,
vation through promotion of recommended agriculture tech- and so on. However, despite the production strength, Indian
nology and a package of practices by addressing the underlying corn yields are significantly below the yields in major corn-
constraints of different agroclimatic subregions. It is under producing countries. There is immense scope for an increment
operation in seven states including Uttar Pradesh, Jharkhand, in Indias corn production by increasing the area under
Bihar, West Bengal, Assam, Orissa, and Chhattisgarh. Rice hybrids, adoption of better genetics, and improved agronomic
contributes to more than 40% of total food grain production practices. Maize is primarily a kharif crop (summer seasons
and is cultivated throughout the country. Rice production in crop) with 85% of the area under cultivation and accounting
India shows a steady upward trend, but it is subjected to wide for 9% of total food grain production. In India the CAGR
year-to-year fluctuations compared to wheat, as a significant (cumulative annual growth rate) of maize production has been
portion of the crop is not irrigated. Indian Basmati rice is 5.5% over the last 10 years from 14 million tonnes in 200405
traditionally grown in Punjab, Haryana, and western Uttar to 23 million tonnes in 201314 (Figure 4). There was a
Pradesh. With the introduction of high-yielding PUSA 1121 decline in the production of maize during 200910 due to
variety, Indias long-grain basmati rice production has been drought conditions in India.
370 GRAINS AROUND THE WORLD | Production and Consumption of Grains: India

10 Area under cultivation Yield 2.9

Area under cultivation (million hectare)

Yield (million tonne/hectare)

8 2.2

6 1.5
5

4
-0

-0

-0

-0

-0

-1

-1

-1

-1

-1
04

05

06

07

08

09

10

11

12

13
20

20

20

20

20

20

20

20

20

20
Years
Figure 5 Yield and area under cultivation of maize.

Area (Hectares) Production (million tonnes) Yield (Kg/Hectare)

3000

2500

2000

1500

1000

500

0
2007-08 2008-09 2009-10 2010-11 2011-12 2012-13
Year
Figure 6 Area, production, and yield of barley in India.

In Figure 5 area under maize cultivation, yield, and CAGR of habits and lifestyle has reduced the demand for barley for
area are mentioned. During the last decade (200405 to human consumption, thereby causing the area in cultivation
201314), CAGR of the area under maize cultivation has of barley to plummet during last few decades. Barley is utilized
increased 2.5%, and cultivated area increased from 7.5 million for beer, food processing, and feed industries apart from direct
hectares in 200405 to 9.4 million hectares in 201314. Factors human consumption in India. Recently, a rise in demand for
such as adaptability to diverse agroclimatic conditions, lower beer in urban India has directly increased the demand for
labor costs, and lowering of the water table in the rice belt of barley malt from Indian beer manufacturing industries.
India have contributed to the increase in acreage of maize. The country is self-sufficient in barley production, although
Productivity of maize has accelerated at a CAGR of 2.9% from production is low, besides exporting 0.10.2 million tonnes of
1.9 million tonnes per hectare in 200405 to 2.5 million tonnes barley annually. In recent years, Indias annual production has
per hectare in 201314. Yield of maize increased by 20% due to been stagnant at 1.21.5 million tonnes, with production in
introduction of single-cross hybrid seeds coupled with adequate 201213 at 1.74 million tonnes against 1.61 million tonnes
rainfall in 200708. In India 85% of maize production and 80% reported last year. Similarly, the area under cultivation has also
of the area under cultivation come from nine states viz. remained stable at 600700 ha, with a per-hectare yield of
Karnataka, Andhra Pradesh, Tamil Nadu, Rajasthan, Maharash- 2508 kg (Figure 6).
tra, Bihar, Uttar Pradesh, Madhya Pradesh, and Gujarat. In India, major barley-growing states are Uttar Pradesh,
Rajasthan, and Madhya Pradesh, with a contribution of 34%,
30%, and 12%, respectively, in total acreage. Compositely,
these states hold about 80% of the whole acreage (Figure 7).
Barley
Rajasthan ranks second in terms of acreage under barley
Barley is the fourth-most-important and most adapted crop of and holds first place in terms of production due to a higher
all cereals. It is used for both feed and food. A shift in the food yield level. Rajasthan accounts for 40% of total production of
 GRAINS AROUND THE WORLD | Production and Consumption of Grains: India 371

protein, which is almost twice the protein in wheat and three

Uttar Pradesh times that of rice. India is the leading producer in the world
16% with 25% of the total share and is also the largest importer and
Rajasthan consumer in the world. Major pulses grown in India include
4% 34% Bengal gram or gram, red gram, lentil, black gram, green gram,
4% Madhya Pradesh moth bean, horse gram, cowpea, and faba bean. Most popular
among these are Bengal gram, red gram, lentil, black gram, and
green gram. Madhya Pradesh, Maharashtra, Uttar Pradesh,
12% Himachal Pradesh
Andhra Pradesh, Karnataka, and Rajasthan are the major
states growing pulses in India. These six states contribute
Haryana 70% of total pulse production and area. The country has
30% achieved a record pulses production of 19.57 million tonnes
Others in 201314 compared to 18.34 million tonnes in the year
201213. Production of all the pulses, as a whole has been
Figure 7 Area under cultivation of barley in various states of India. increasing year after year except for a downturn in 201112.
The production of black gram only has been estimated to
decline from 1.9 million tonnes in 201213 to 1.5 million
Rajasthan tonnes in 201314.
9%
5% Uttar Pradesh
6% Consumption of Food Grains
40%
Madhya Pradesh
9%
In India the rate of growth of population and food grains
production for last five decades are illustrated in Figure 9. It
Haryana has been observed that the population growth rate has seen a
downfall from 2.24% to 1.03%. The overall rate of production
Punjab of food grains has also declined, with a rise only during the
31% period from 198191. During 200405 total cereal consump-
tion showed a rise of about 100 g per person per month in
Others
rural areas and a fall of 350 g per person per month in urban
Figure 8 Barley productions in Indian states. areas. The per capita consumption trend of grains such as
sorghum, pearl millet, and maize has been higher in the rural
sector as compared to a declining trend in consumption of
barley followed by Uttar Pradesh (31%), Madhya Pradesh
major grains like rice and wheat. The cereals other than rice
(9%), and Haryana (6%; Figure 8).
and wheat contributed 1 kg per person per month in rural
India and 0.8 kg per person per month in urban India in
Coarse Cereals 201112.
The consumption of all food grains per person in the rural
Crops like sorghum, millets, ragi, and other small millets (kudo, areas is projected to decline from 15.3 kg per month in 2000 to
kutiki, sanwa, and foxtail) termed as coarse cereals have been the 13.8 kg per month by 2050 and to only slightly decrease from
primary components of the food basket of rural India. Rain-fed 11.8 to 11.6 kg per month in the urban areas. It has been
regions of India like Karnataka, Maharashtra, Tamil Nadu, Mad- estimated that due to rapid urbanization, per capita consump-
hya Pradesh, Rajasthan, and Gujarat are best suited for coarse tion of food grain in India will decrease from 14.4 to 12.7 kg
cereal crops, and coarse cereals are therefore cultivated predom- per month over the next 50 years. However, total food grain
inantly in these areas. A downturn has been witnessed in the area demand is projected to increase from 16.7 to 19.9 kg per
covered under coarse cereals cultivation from 29.03 million month over the next 50 years due to increment in the demand
hectares in 200405 to 26.42 million hectares in 201112. of feed grain. The total grain demand will increase from 201
Significant increases in the yield of coarse cereals has been wit- million tonnes in 2000 to about 291 and 377 million tonnes
nessed from 1153 kg ha 1 in 200405 to 1591 kg ha 1 in by 2025 and 2050, respectively.
201112. Production of coarse cereals in 200405 was 33.46 The major diet of the Indian people is vegetarian, and food
million tonnes, which increased to a maximal level of 43.40 grains are considered the main components of their diet,
million tonnes in 201011 but decreased slightly to 42.04 mil- although some variation in diet is found across different parts
lion tonnes in 201112 mainly because of a shift in area of of India. Wheat is the principal choice of food grain in the
cultivation to other competing crops. north followed by rice. Wheat comprises half of the total grain
consumption in the west followed equally by rice and other
coarse cereals together. Wheat is the staple food in India mostly
Pulses
consumed in the form of homemade chapattis or rotis (unlea-
Pulses complement the staple cereal in the diet as a gifted vened flat bread) using atta (whole wheat flour). Wheat is also
source of proteins, essential amino acids, vitamins, and min- consumed as bakery products like pan breads, cakes, biscuits,
erals among the vegetarians in India. They contain 2224% cookies, and crackers.
372 GRAINS AROUND THE WORLD | Production and Consumption of Grains: India
Population
Growth rate (Percentage)
4
0
1961-71 1971-81
Figure 9 Growth rate of population and food grain production in India.
Rice is consumed as the main food grain in the south and
the east followed by wheat in the east and coarse cereals in the
south. Rice consumption in 201314 was estimated at 95.0
million tonnes compared to 93.5 million tonnes in 201213, a
modest increase of 1.5%, almost equal to the population
growth rate of 1.4% per annum.
Monthly rice consumption in India declined to the extent of
400 g in rural and 220 g in urban areas from 6.38 to 5.98 kg to
4.71 to 4.49 kg during, respectively, the period from 200405
to 201112 in rural and urban areas, respectively. On the
contrary, monthly consumption of wheat showed a slight
increase of about 100 g per person in rural areas and a fall of
350 g in urban areas during the same period.
Direct human consumption of maize has been reduced
over the years. In 201213, consumption of maize in the
form of human food was only 20%, and it is estimated to
decline further due to rising per capita income and
commercialization, which has made cereals like wheat and
rice more preferred and affordable. Maize is the basic raw
material utilized for starch manufacturing and increasing com-
mercial demand from poultry and starch industries leading to
higher farmer realization. The remarkable portion, that is, 47%
of total maize production is used for poultry feed processing
because it possesses good nutritional value. Out of the total
production of maize, 12% is for starch manufacturing, 14% for
cattle feed, and 7% for brewery and food processing
(Figure 10). Maize contains 6065% starch content and
hence cannot be easily substituted by other commodities for
starch manufacturing.
Barley was the staple food for many countries until wheat
supplanted it, resulting in the deceleration of human con-
sumption of barley with time. A wide variety of barley products
known to be suitable for human consumption includes por-
ridge, muesli, cookies, barley flakes, muffins, breads, and
pasta. Barley, being a good source of protein, is used as a feed
for livestock. Barley is used as animal feed in the form of rolled,
ground, flaked, or pelleted grain. Barley is also used in alco-
holic beverages like beer and wine in the form of malt. Apart
from this, malted barley is also used as flavors, sweeteners,
malt extracts, malt flours, and so on. In India total malt pro-
duction is around 3.5 lakh tonnes, but the demand of beer and
Foodgrain Production
1981-91 1991-01 2001-11
Years
7% Poultry
12%
Direct Consumption
47% Cattle Feed
14%
Starch manufacturing
20%
Food Processing
Figure 10 Utilization of maize in India.
malt is increasing at the rate of 10% annually; to meet this
demand, the area and production of the barley needs to be
increased.
Economic growth has continued to fuel increased pulse
consumption in recent years. Consumption in 201415 is
forecasted higher at 22.5 million tonnes compared to 21.5
million tonnes last year on continued strong domestic demand
for the rising Indian population. Monthly consumption of
pulses and their products increased from 705 to 783 g in the
rural sector and from 824 to 901 g in the urban sector for this
period. The growth in the pulses consumption was attributed
mainly to the increase in the consumption of split gram, whole
gram, peas, and besan (Bengal gram flour) to the extent of 69 g
in the rural sector and 57 g in the urban sector. The nongrain
food crops and animal products contributed 27% and 8%,
respectively, of the total caloric requirement. The meat con-
sumption in India is very low due to cultural and religious
reasons, and most of the calories from animal products are
obtained from milk and milk products.
Future Consumption Trend
The emerging food consumption trends in India have been
illustrated by many authors. They have claimed that in recent
decades there has been a clear transition from grain
 GRAINS AROUND THE WORLD | Production and Consumption of Grains: India
consumption to nongrain food and animal product consump-
tion. Rath suggested that the per capita grain consumption will
further decrease due to the reduction in physical labor require-
ment in rural areas due to the adoption of more mechanized
and modernized agriculture. Furthermore, improving medical
facilities and infrastructure also contribute to the reduction in
energy requirements and thus less cereal consumption. It has
been suggested that a reduction in the intake of food grains on
this account should not be taken as deterioration in human
welfare.
Sustained economic growth and changing lifestyles are
accounting for a significant shift in Indian food culture that is
inclined toward high-value horticultural and animal products
and away from staple food grains. Rising per capita income and
urbanization are leading to a dietary transition primarily from
cereals with low-price calorie foods (foods such as wheat and
rice with low price and rich in carbohydrates) toward high-
price calorie foods like livestock products and processed or
ready-to-eat products. The transitions are also attributed to
farming patterns, as farmers are adopting new technologies
that have resulted in an increase in farm productivity.
In accordance with Engels law, as the earnings of people
rise, the cheaper source of energy for living, which comes from
the staple food consumption, is likely to drop. A decline in
cereal consumption is substituted with the increased consump-
tion of high-value commodities. Over time the shifting of
cereal consumption pattern will be prominent across all house-
holds in different expenditure groups. The devolution in cereal
consumption can also be credited to the diversification of food
production, easy access of high-value commodities, and
switching tastes and preferences. This agrees with the argu-
ments by Kumar and Mathur that the demand for food is not
only influenced by the income of a person but also by differ-
ences in urban and rural lifestyles, the development of more
advanced marketing systems, and occupational changes are
closely linked with increasing per capita income. Tastes and
preferences are also moving toward high value commodities.
With the availability of cash money and high-value commod-
ities in the food basket, the decline in calories by reduced cereal
consumption is partly compensated for by intake of highly
nutritive and vitamin-rich food products. Some other studies
have also attributed the change in per capita consumption of
cereals to economic growth, rise in per capita income, urban-
ization and preferences, market integration, and many more as
the dominating factors.
Exercise for Revision
How has Indian agriculture been successful in changing its
face postindependence?
In what ways has the Indian government contributed to the
growth of agriculture in India?
What factors govern the changing consumption patterns of
food grains in India, and how is it changing?
What is the total production of different grains in India, and
which are the major grain-producing states?
373
Exercises for Readers to Explore the Topic Further
Why are Indias food grain production and productivity still
lagging behind?
What would be the implications of economic development
and urbanization on the food grain consumption patterns
of the Indian population?
What would be the effects of switching consumption pat-
terns on the future production estimates of grains?
See also: Food Grains and the Consumer: Consumer Trends in
Grain Consumption; Cultural Differences in Processing and
Consumption; Grains Around the World: Grain Production and
Consumption: Overview; The Basics: The Grain Chain: The Route
from Genes to Grain-Based Products; The Grain Crops: An Overview;
The Cereal Grains: An Overview of the Family of Cereal Grains
Prominent in World Agriculture; Barley: An Overview of a Versatile
Cereal Grain with Many Food and Feed Uses; Maize: Overview; Rice:
Overview; Wheat: An Overview of the Grain That Provides Our Daily
Bread.
Further Reading
Amarasinghe UA, Shah T, and Singh OP (2007) Changing consumption patterns:
Implications on food and water demand in India. Colombo, Sri Lanka: International
Water Management Institute, 43 p. (IWMI Research Report 119).
Annual Report, 201112. Department of Agricultural Research (DARE) & Education/
Indian Council of Agricultural Research (ICAR).
Bansil PS (1999) Demand for foodgrains by 2020 AD. New Delhi: Observer Research
Foundation.
Directorate of Economics and Statistics, 1213. Department of Agriculture and
Cooperation, Ministry of Agriculture, Government of India, New Delhi.
Kumar P and Mathur VC (1996) Structural changes in demand for food in India. Indian
Journal of Agricultural Economics 51(4): 664673.
Kumar P and Mathur VC (1997) Agriculture in future: Demand and supply prospective.
In: Desai BM (ed.) Agricultural Development Paradigm for the Ninth Plan Under New
Economic Development. New Delhi, India: Oxford and IBH Publishing.
Mittal S (2007) What affect changes in cereal consumption? Economic and Political
Weekly: 444447, February.
National Sample Survey (NSS)-68th round July 2011-June 2012. Household
consumption of various goods and services in India, Government of India.
Radhakrishna R and Ravi C (1990) Food Demand Projections for India. Hyderabad:
Centre for Economics and Social Studies.
Radhakrishna R and Ravi C (1992) Effects of growth, relative price and performances of
food and nutrition. Indian Economic Review 27: 303323.
Rao CHH (2000) Declining demand for foodgrains in rural India: Census and
Implications. Economic and Political Weekly, January 22.
Rath N (2003) Linking of rivers: Some elementary arithmetic. Economic and Political
Weekly: 30323033, July.
Singh SK (2014) Grain and feed annual, India. Global Agricultural Information Network
(GAIN) report-IN4005.
The Economic Survey 201415, Agricultural and Processed Food Products Export
Development Authority (APEDA).
The Tribune, Chandigarh, February 21, 2014. Sale of beer on the rise, scientists for
contract farming of barley.
Relevant Websites
faostat.fao.org FAOSTAT, data.
 Grain Production and Consumption: China and South-East Asia
H Corke and YZ Cai, The University of Hong Kong, Hong Kong, Peoples Republic of China
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition volume 2, pp 7786, 2004, Elsevier Ltd.
Topic Highlights
Asia is characterized by high and increasing populations,
and limited and declining areas of arable land.
Asian countries have to balance a desire for food self-
sufficiency requiring great agrotechnical and social resource
commitments, with an openness to import staple grains
which carries political risks such threat of sanctions or
boycott.
The Green Revolution of the 1960s, greatly increasing rice
and wheat yields in Asia, and transforming India into a
grain exporter, led to greater optimism about longterm
prospects for food production.
Chinas production and consumption, and Chinas increas-
ing demand for meat and other animal products, will con-
tinue to be a major driver of global grain production and
marketing.
Learning Objectives
To understand the scale of Asian grain production, the
population and land pressures, and the necessity for most
Asian countries to pursue substantial self-sufficiency in
grain production.
To understand the significance of the Green Revolution for
Asian grain production.
To understand the relationship between demand for meat
and demand for grain.
Without agriculture, there is no stability, without grain, there is
chaos.
Deng Xiaoping
Introduction
Several factors make Asian grain production and consumption
key to understanding the global balance of grain exports,
imports, and food sufficiency. Asia has the worlds two most
populous countries, China with  1.2 billion people, and India
with  1 billion and on track to exceed China in population
within 2030 years. Asia is the dominant producer of rice, yet
an insignificant amount enters world grain trade. Asia is a major
net importer of wheat and soybeans, thus influencing decisions
about quality priorities in exporting countries. Asian countries
have tended to aim for grain self-sufficiency for reasons of
agrarian stability, foreign currency conservation, and indepen-
dence from political restrictions on their imports from major
western producers. Major successes in grain production over
recent decades have increased confidence in regional capability
374 Encyclopedia of Food Grains, Second Edition
to seek continuing technological solutions to create ever-
increasing productivity. This confidence has been dented by
stalling yield gains, especially in overexploited rice soils, and
by industrialization and urbanization, which absorb agricul-
tural land and compete for water resources. This article will
discuss these issues, focusing on three major themes: (1) statis-
tical patterns of grain production, import, and export; (2) the
impact and aftermath of the Green Revolution in wheat and rice
production; and (3) the role of China as a consuming giant in
an era where small-scale grain production makes less economic
sense, and where increasing prosperity drives demand for grain-
intensive animal product consumption. Asia is a diverse region
with many countries of widely differing land area, population,
and specific problems. These will not be discussed separately,
but the main issues will be presented by discussion of the few of
the biggest countries.
The General Problem
Although rice is the staple crop for half the population of the
world, only 2% of total production is traded on international
markets. The principle of national production for domestic
consumption has driven most Asian countries to aim primarily
for self-sufficiency (although some countries such as Thailand
and India are rice exporters). In many countries, such as Phil-
ippines, self-sufficiency is difficult to achieve. Populations
are increasing; people are increasingly moving off the land to
the cities; water supply is not sufficient to produce the possible
three harvests per year on the rice paddies; every flat arable piece
of land is already in production, so major land-use changes
tend to be those taking land out of production. In the Philip-
pines and elsewhere, rice has in many areas been grown con-
tinuously in the same fields for centuries. It had long been
supposed that atmospheric and water-borne mineral and trace
element additions to the soil are sufficient to allow indefinite
maintenance of soil fertility, requiring only macronutrients and
water for each new crop. But results of long-term field experi-
ments analyzed at the International Rice Research Institute
(IRRI) near Manila showed that across Asia, for irrigated rice
systems, the soil base has been depleted of nutrients by contin-
uous monoculture. The result of this is that the same variety
responds less to the same level of added nitrogen. In other
words, improved varieties and production methods are needed
simply to maintain current yield levels. In future, expanded use
of hybrid rice varieties using Chinese technology may increase
yield per unit area but also further increase mineral depletion
and biomass removal from the field. Philippines has achieved
approximate self-sufficiency in rice in 2004 but this may be
transient; demand will continue to increase, and energy-
intensive inputs will still be required. Farmers who produce
rice on small pieces (typically 2 ha) of rented land will continue
http://dx.doi.org/10.1016/B978-0-12-394437-5.00045-0
 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia 375

to seek lives with less physical hardship and better incomes populous nation in the not too distant future. The limited land
elsewhere. supply (an index of population density) in China is going to be
In a 1996 interview, K. Lampe, then Director General of matched across Asia as a whole as the population of the rest of
IRRI, summarized the problems of developing tropical Asian Asia continues to increase (Figure 1). Although the population
economies such as Philippines. density of India is greater than China, the supply of arable land is
also greater (Figure 2). Widely quoted figures are that China has
22% of the worlds population but only 7% of the worlds arable
In 30 years from now we will have for example in the Philippines land. This creates a stark problem how to feed  5 billion
120 million people, 60 million more than today. How to feed them, Asians in 25 years time without increased supply of high-quality
nobody seems to care. Over the last 30 years rice has had to increase land (indeed, with high-quality land continuing to be lost to
to such an extent that 600 million people can eat rice today who
otherwise would not be able to. Another 30 years and we are
urbanization and industrialization). Six Asian countries had
predicting a world population of more than 8 billion people. More populations of 100 million or more in 2001 (Figure 2). Their
than half of them will be rice eaters. A prediction from United agricultural populations accounted for 4366% of their total
Nations data is that about 400 million people will move over the population, except for Japan, which had only 3.6% of its people
next 10 years from rural areas in Asia to the big cities. What will
engaged in agriculture. Japan, as a highly advanced economy
happen is that the belt of poverty around these mega-urban con-
glomerations will become most probably unbearable. And what we which is already committed to food import rather than food self-
will face is most probably social unrest of unprecedented magni-
tude. Our task is to grow more rice on less land with less fertilizer,
less pesticides, and less labor, because producing rice is one of the China
most tedious tasks for a farmer.
India
Kazakhstan
Saudi Arabia
Indonesia
Iran
Production Statistics Mongolia
Pakistan
Using the Food and Agriculture Organization of the United Turkey
Nations (FAO) definition of Asia, the population of Asia has Myanmar
Others
more than doubled in the past 40 years (Figure 1), and is
dominated by its two most populous countries (China and 0 20 40 60 80 100
India). The population of China is more or less under control, Land area (10 Mha)
such that India is likely to overtake it as the worlds most
India
China
Turkey
6000
Kazakhstan
5000 Pakistan
Population (million)

Asia Indonesia
4000
Thailand
3000 Iran
Myanmar
2000 Bangladesh
China
1000 Others
India
0 40 80 120 160 200
0
Arable land (Mha)
1950 1965 1980 1995 2010 2025 2040 2055
Years China
1.9 India
Indonesia
Land area (ha per capita)

1.7
Pakistan
1.5 Aisa Bangladesh
1.3 Japan
1.1 Vietnam
0.9 Philippines
0.7 China Iran total pop. (M)
0.5 Turkey
agric. pop. (M)
India Others
0.3
0.1 0 200 400 600 800 1000 1200 1400
1950 1965 1980 1995 2010 2025 2040 2055 Population (million)
Years
Figure 2 Land area, arable land, and total and agricultural populations
Figure 1 Estimated population (19502001) and land area per capita of top ten population countries in Asia in 2001. (Data from FAOSTAT
(19612000) of all Asia, China, and India, with projections till 2050. Database).
376 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia

sufficiency, has substantially different problems from other
Asian countries. It focuses on managing the quality and suitabil-
ity of its imports, and on maintaining a domestic agricultural
sector (e.g., a meat industry using imported grain), which com-
petes poorly on price but which is politically and socially
important.
China produces 41% of Asian grain on 26% of the area
under cultivation; India produces 22% on 30% of the area
(Figure 3). This reflects a wider adoption of intensive cultural
practices and some favorable environmental factors in China.
Indonesia, another highly populous country, also has greater
production per unit area than the average for Asia, whereas
Thailand shows below-average production. These figures can
be used to indicate possible areas for greater technology adop-
tion and hence increased production using existing land under
cultivation.
How is Asias one billion tonnes (Gt) annual grain produc-
tion distributed across crops? Asia is well-known for rice pro-
duction, and for the widespread dietary cultures based on rice
consumption. Indeed, more than half of Asian grain produc-
tion is rice (Table 1, Figure 4) with 523 million tonnes (Mt) in
2002. Other than for India, Asia is less well known as a wheat
producer, yet more than 44% (Table 2) of global wheat pro-
duction is from Asia, and China is by far the largest wheat
producer in the world. Maize production at 166 Mt, again
predominantly in China, is substantial. China ranks second
to the US in maize production. It is interesting that the center
of origin of maize is in America (Mexico) but it has become
globally widespread and successful, including in Asia. How-
ever, soybean, a crop originating in Asia, is more commercially
significant in the US than in Asia. Sorghum and millets are
significant crops in arid parts of Asia, typically areas of China
and India too dry for maize production. Many local traditional
uses of these cereals have developed, for example typical use of
sorghum in China is in distilled alcoholic beverage production.
Buckwheat also has a significance beyond its relatively small
production, with uses in traditional soba (Japanese buck-
wheat noodles) and some Chinese traditional foods such as
noodles, vermicelli, and fermented vinegars. Barley is a sub-
stantial crop, fourth after maize, partly reflecting Chinas rise to
rank first in the world for beer production.
Asias grain imports in 2001 were worth about US$ 25
billion, and exports about $8 billion (Table 2). Imports were
dominated by wheat, maize, and soybean, and exports by rice.
Industrial, processed food, and feed uses predominate for
maize and soybean. The unique properties of wheat flour
make it impossible to substitute with any other grain. Strong
demand for wheat supplies exists for traditional uses (e.g.,
noodles and steamed bread in northern China and other tem-
perate Asian countries). Economic development also tends to

India 29.7%
China 26.4%
Indonesia 4.9%
Turkey 4.6%
Pakistan 3.9%
Bangladesh 3.9%
Thailand 3.7%
Vietnam 2.7%
Myanmar 2.2%
Philippines 2.1%
Others 14.1%
0 20 40 60 80 100
Harvested area (Mha)

China 40.8%
India 21.7%
Indonesia 6.2%
Bangladesh 4.0%
Vietnam 3.7%
Turkey 3.2%
Thailand 3.1%
Pakistan 2.7%
Myanmar 2.3%
Philippines 1.8%
Others 10.5%
0 90 180 270 360 450
Grain production (Mt)
Figure 3 Harvested area and grain production of top ten grain-producing countries in Asia in 2002. Data from FAOSTAT Database. Percentage (%) is
based on total harvested area or total grain production in Asia.
 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia 377

Table 1 Area, production, and yield of grains (cereals, soybeans, and pulses) in Asia (2002)a

Area harvested Production

Crop Mha % of Asia total Mt % of Asia total % of world total Yield (kg ha 1) Countries (number)

Cereals, total 307 100 986 100 48.6 3209 47

Paddy rice 131 42.6 523 53.1 90.8 3998 30
Wheat 96.1 31.3 253 25.6 44.1 2629 37
Maize 42.7 13.9 166 16.8 27.5 3884 39
Barley 11.8 3.8 19.4 2.0 14.7 1645 33
Millet 11.4 3.7 9.1 0.9 39.0 803 20
Sorghum 11.4 3.7 11.0 1.1 20.2 959 20
Buckwheat 0.9 0.3 1.4 0.1 65.2 1451 6
Rye 0.7 0.2 1.1 0.1 5.2 1467 11
Oats 0.7 0.2 1.1 0.1 4.3 1543 15
Triticale 0.5 0.2 1.2 0.1 10.9 2320 1
Others 0.2 0.1 0.2 0.0 2.6 2641 11
Soybeans 17.1 23.7 13.2 1385 25
Pulses, total 36.3 25.6 46.4 706 40
a
Data from FAOSTAT Database.

250
450 China
China Bangladesh
Bangladesh
400 India Indonesia
India
Total grain production (t)

200 Thailand Vietnam

350 Indonesia
Rice production (t)

300 150
250
200 100
150
100 50
50
0 0
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005
Years Years

140
120 China India China India
Iran Kazakhstan 120 Indonesia Philippines
100 Pakistan Turkey Thailand Turkey
100
Wheat production (t)

Maize production (t)

80
80

60 60

40 40

20 20

0 0
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005
Years Years

Figure 4 Trends of yield production in major Asian countries producing principal cereal crops from 1961 to 2002. (Data from FAOSTAT Database).

increase demand for wheat products (e.g., baked bread) at the
expense of rice consumption. Thus, many tropical Asian coun-
tries that do not produce any wheat require substantial imports
(e.g., Philippines, Indonesia, Malaysia). Country data (Table 3)
show that Japan is the largest net importer of grain in Asia. Its
demand is likely to be stable, due to constant (or slightly
declining) population and its advanced economic status
(far past the rapid growth stage) and therefore stable dietary
patterns. China was a major importer but also a significant
exporter (in 2001). But its importexport pattern is highly
variable, as slight percentage changes in its large production
base, will cause dramatic changes in its import needs and
export capacity (Figure 7). India was a net exporter in 2001,
and should be able to maintain this capability in the short
378 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia

Table 2 Quantity and value of grain imports and exports to/from Asia in 2001

Crop Imports qty (Mt) Imports value (1000 US$) Exports qty (Mt) Exports value (1000 US$)

Cereals 106 478 996 16 732 160 38 394 615 7 057 804
Maize 40 913 789 5 076 869 7 013 319 759 271
Wheat 40 238 186 6 760 921 7 906 977 896 782
Barley 10 117 996 1 490 596 436 870 40 048
Rice 9 426 314 2 623 935 19 798 418 4 902 134
Sorghum 2 015 717 233 845 20 587 3061
Rye 381 226 46 196 15 983 2065
Oats 112 604 20 100 5597 701
Buckwheat 104 668 25 227 104 491 20 365
Millet 65 854 15 188 45 735 9099
Triticale 251 74 25 528 5653
Others 3 102 391 439 209 3 021 110 418 625
Soybean 28 045 849 6 025 734 366 951 109 096
Pulses 4 463 201 1 468 603 2 636 496 1 065 219

Table 3 Top ten Asian countries of grain (cereal and soybean) imports and exports in 2001

Import quantity Export quantity Net import quantity

Top ten import Percentage Top ten export Percentage Top ten net import Percentage
countries Mt (%) countries Mt (%) countries Mt (%)

Asia, total 134.5 100.0 Asia, total 38.8 100.0 Asia, total 95.8 100.0
1. Japan 31.1 23.1 1. China 9.3 23.9 1. Japan 30.1 31.4
2. China 26.3 19.6 2. Thailand 8.2 21.2 2. China 17.1 17.8
3. Korea 13.7 10.2 3. India 5.4 14.0 3. Korea 13.7 14.3
4. Iran 10.4 7.7 4. Vietnam 3.8 9.8 4. Iran 10.4 10.8
5. Indonesia 5.9 4.4 5. Kazakhstan 3.6 9.2 5. Indonesia 5.8 6.1
6. Saudi Arabia 5.1 3.8 6. Pakistan 3.4 8.6 6. Saudi Arabia 5.1 5.4
7. Malaysia 4.4 3.3 7. Turkey 1.6 4.0 7. Malaysia 4.3 4.4
8. Philippines 4.2 3.1 8. Myanmar 1.0 2.7 8. Philippines 4.2 4.4
9. Iraq 4.0 2.9 9. Japan 1.0 2.6 9. Iraq 3.9 4.1
10. Israel 3.6 2.6 10. United Arab 0.5 1.3 10. Israel 3.6 3.7
Emirates
Others 25.8 19.2 Others 1.1 2.8 Others 2.3 2.4

term. Other countries have adopted a policy that will see

30
sustained dependence on imports, some not having any eco-
Change during 19922002 (%)

nomic reason or climatic suitability to pursue self-sufficiency 20 harvested area

grain production
(e.g., Saudi Arabia). Sustained import markets are the easiest 10
for exporters to deal with they can develop relationships with 0
the buyers to understand quality needs and develop long-term
10
policies to meet these needs. Erratic, but large purchases, on
international markets are more difficult to handle efficiently. 20
Total cereal production has risen fairly steadily across Asia as 30
a whole since 1960 (Figure 4) (despite Chinese production in 40
the last 5 years, although high, has been more variable than in
50
the past). This reflects improved varieties and agronomic prac-
tices (under the Green Revolution, discussed below), but also 60
, p al

W y

M t
ze

So ley

w t
at

Tr s

O le
s

expansion of cultivated area under the multipronged

a

ille
d

at

er
Ry
ce ot

ca
he

hu

he
ai
ad

th
M
Ba
Ri s, t

iti
rg

approaches adopted to cope with vast increases in demand.

ck
l
ea

Bu

The situation has become rather less stable in more recent

er
C

years (Figures 4 and 5), with substantial drops in production Cereal crops in Asia
of cereal grains other than the big three, namely, rice, Figure 5 Change in Asian grain production and harvested area of
wheat, and maize. In fact, between 1992 and 2002 total pro- total cereals by crop over the past decade (19922002). (Data from
duction (output) increased only  6%, whereas area harvested FAOSTAT Database).
 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia
decreased  8%. Although this reflects a healthy improvement
in yield per unit area, it underscores worrying trends. One is
the diversion of agricultural land to noncereal uses, offering
higher returns to farmers, and another is the steady loss of
high-quality agricultural land. In many places, such as in parts
of China, production of grain on small family-based farm
units is no longer economically attractive. Other agricultural
activities vegetables, meat production such as ducks, or even
flower production may be preferable. In some cases, land may
be left idle while the landholder pursues nonagricultural
business activities. These shifts are often beneficial in raising
the living standards of rural families. But they also show that
more or less open markets, keeping grain prices near world
levels, are certainly incompatible with full self-sufficiency in
grain.
The Green Revolution
India has all the food it needs, but half of it is currently being
eaten by rats, according to a western aid official quoted in a
Financial Times report of December 2002. In the three years up
to 2002, Indias reserve of rice and wheat increased from 20 to
60 Mt, relative to an annual production of 220 Mt. Half of
Indian children are underweight, and many are malnourished.
India has seemingly solved one problem the technological
problem for which the solution began with the Green
Revolution only to be left with a perhaps more intractable
problem, that of creating an economic system that distributes
food equitably and without undue wastage.
Development of dwarf wheat that could sustain high fertilizer
inputs and thus produce high grain yields without lodging began
at International Center for the Improvement of Maize and Wheat
(CIMMYT) in Mexico in the early 1960s. At that time, Indias
wheat production was low ( 10 Mt per year) and static, showing
only random seasonal fluctuations from year to year. Like most
low-income, high-population countries, population growth was
much more rapid than growth in food production, and a social
catastrophe seemed imminent. The dwarf wheat from Mexico
was introduced to India, through a system of demonstration
fields and a supporting package of technology that emphasized
to farmers the uniqueness of the new varieties, and the necessity
of growing them with the high inputs of water, fertilizer, and
other agricultural chemicals. From the mid-1960s, wheat pro-
duction in India began to increase rapidly, outstripping growth
in population and leading to the present day problem of
distribution of excess production. Over a 20-year period
(196180), overall food production increased 3.6% per year in
Asia as a whole, from 1981 to 2000 (sometimes referred to as
Late Green Revolution), it increased 2.1% per year. Thus, we
should consider the adoption of high-yielding, high-input wheat
varieties as the beginning of an agricultural transformation, and
not as an abrupt one-time revolution. Present-day technologies
may be seen as contributors to the same continuum of progress.
If production increases in the range of 23% a year, clearly
limiting the population growth to less than this is the key to
increased production per person.
Shortly after the dwarf wheat varieties came into use, a
parallel development started with dwarf rice varieties, distrib-
uted from IRRI Philippines. G. S. Khush, the IRRI plant breeder
379
responsible for much of development of these rice varieties,
summarized their biological features:
reduction in plant height (improved harvest index) and
higher biomass,
photoperiod insensitivity (can grow any time of year) and
short growth duration (110 days),
increased yield stability through disease and insect resis-
tance (biotic stresses),
tolerance to adverse soil conditions (abiotic stresses),
eating quality, and
comprehensive agronomic knowledge on crop manage-
ment, e.g., nutrient requirements, water management,
mechanization techniques, and equipment.
These technical inputs alone were not enough. As with wheat
in India, production conditions had to be improved. The main
factors associated with this are:
Increased provision of irrigation facilities,
Availability of inorganic fertilizers, and
Supportive government policies
The Green Revolution is a high-input system. There is no free
lunch or nonenergy intensive way to produce such dramatic
increases in production. Central to this is farmer knowledge
and participation in a system with higher risks (e.g., use of
credit to finance the required inputs) but better rewards
(higher economic returns).
The Crisis
Grain Supply in China
Chinese tradition and the political leadership have a clear
understanding of the importance of grain production, from
the proverb Of all things food is the foremost necessity of
the people to the comment by Deng Xiaoping Without agri-
culture, there is no stability, without grain, there is chaos. This
has an impact even in relation to Hong Kong: As long as the
overall political situation in China is stable and the economy
develops further, we will stand by our agreements and princi-
ples agreed with Britain Qiao Shi (quoted 8 January 1995).
This was in relation to the agreed transfer of sovereignty of
Hong Kong from the United Kingdom to China from 1997,
where substantial autonomy would be granted to Hong Kong
(as a Special Administrative Region) for a period of 50 years.
Pragmatically speaking, if a government cannot feed its popu-
lation, it cannot maintain the credibility to stay in power and
carry out its normal obligations. This issue is all the more
pressing in China with its large population, limited area of
arable land, and recent memory of the disastrous famine of
the early 1960s, which caused the deaths of over 30 million
people.
Money, Meat, and Grain
The book Who Will Feed China? by US environmentalist L.
R. Brown, (1995), was perhaps the most influential early study
in drawing the attention of western policy-makers to a fairly
obvious problem, i.e., grain trade and capacity for global
380 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia
increases in grain production are limited, economic prosperity
increases demand, industrialization reduces production, and
the huge population of China makes it the key country where
these issues converge. Industrialization and urbanization have
many effects on agriculture:
1. Loss of agricultural land to urbanization takes out the pro-
ductive farmland, and of course that nearest to the centers
of population;
2. Increased use of motor vehicles particularly new oppor-
tunities for personal ownership of cars, creates need for
expanded highways, roads, and parking systems, which
also absorb agricultural land;
3. Competing demands for water reduces that available for
agriculture and increases the relative cost;
4. Bringing inferior land under cultivation on the peripheries
of existing arable land use results in lower average yields,
and increased susceptibility to erosion, salinization, and
other environmental problems;
5. The movement of people from rural to urban areas in
search of better paying jobs reduces agricultural production
capacity; and
6. Intensive land use requires ever-greater input of agricultural
chemicals and other energy-based inputs, simply to main-
tain production.
Browns book had pointed out that the entire world cannot
reasonably aspire to the US standard of living and we cannot
afford to keep adding 90 million people a year indefinitely.
In food production terms, the difference between the US
and other countries is the total per-person demand for grain.
Consumption of animal products with the animals consum-
ing grain-based feed makes US per capita grain consump-
tion  800 kg per year. Even Italy, an affluent country
with many cultural affinities with the US, has a consumption
of only 400 kg per person (and a healthier diet with less
obesity). Prediction of Chinas future grain consumption
currently  330 kg per person is the key to preparing to meet
future demand. An earlier study Grain in China published by
the Department of Foreign Affairs and Trade, Australia (1992),
had set out in some detail the technical issues related to this. In
general, increased prosperity does not lead to large increases in
the direct consumption of grain, assuming basic adequacy in
dietary energy intake. However, increases in disposable
incomes do lead to marked changes in dietary composition,
first in the direction of more diversity, and particularly more
meat products (and dairy, eggs, fish, etc.). Further increases in
incomes will increase consumption of processed foods, meals
prepared outside the home, and other convenience options.
Taiwan, with a Chinese population of similar dietary customs,
can be used as proxy for studying future dietary changes with
increased incomes in the Chinese mainland. As mainland
China reaches the purchasing-power-adjusted GDP per capita
prevailing in Taiwan in the past, the demand for particular food
products should also match Taiwans at that stage of develop-
ment. Using data up to 1990, Figure 6 shows that an increase in
mainland Chinas GDP per capita from $1000 to $2000 would
be expected to increase demand for meat by  10 kg per person
per year. Such figures are the basis for concern about Chinas
future needs. An increase in demand from 2003 onwards of a
30
25
Meat (kg)
20
15
Taiwan
10
China
5
200 500 800 1100 1400 1700 2000 2300
GDP (1987 US$)
Figure 6 Per capita GDP and meat (pork, beef, and mutton)
consumption in China (197790) and Taiwan (195275). GDP for
China is increased threefold to adjust for purchasing power.
(Adapted from Garnaut R and Ma G (1992) Grain in China, 151 pp.
Canberra: East Asia Analytical Unit, Department of Foreign Affairs and
Trade, Commonwealth of Australia).
further 10 kg would represent a need for perhaps 40 Mt of grain
(depending on feed conversion ratios and the proportion of
grain used in the animals diet). Naturally, one way to proceed
is to greatly increase meat imports, with the US and Brazil
representing source countries with further potential. However,
this, to some extent, just shifts the burden of grain production
elsewhere (away from Asia), and would deprive farmers in
China of added income if they were simply to produce low-
cost commodity grain (at low production efficiency) while
imports satisfied needs for higher value products.
For Asias two biggest countries, we can summarize that
Chinas main driver of demand, for grain, is shifting food
consumption patterns (toward meat), whereas Indias main
driver of demand is increasing population.
Food Price Rises May Herald a World Crisis
This November 2003 news item published in the South China
Morning Post, Hong Kong, quoted Lester Brown Sudden
food price rises on the [Chinese] mainland could be the
sign of a coming world food crisis. I view these as the warning
tremors before the earthquake. Four consecutive years of
world grain harvest decline, and historically low world
grain stocks since records were kept, could lead to soaring
grain prices worldwide. In 2003, wheat prices in northeastern
China rose 32%, maize prices by 100%, and rice by 13%.
China faced a 40 Mt shortfall in 2003, and the rest of the
world, 56 Mt.
In an era of global grain trade (Figure 7) and heightened
expectations of diminishing barriers to trade, all consumers
(nations) will increasingly compete on price for the basic
staples necessary for survival. Virtually all major grain pro-
ducers still use price-distorting mechanisms such as farm sub-
sidies (in some countries, payments not to produce grain; in
others, artificially high domestic prices, usually coupled with
tariffs and quotas to restrict trade).
More than just a hill of beans headed a Washington Post
article of November 2003 (published in The Standard, Hong
Kong). This illustrated the complex relationship among farmer
 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia
160
Quantity of imports and exports (Mt) Imports
140
120
100
80
60
40
20
0
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005
(a)
35
Imports
Quantity of imports and exports (Mt)
30
25
20
15
10
5
0
1960 1965 1970 1975 1980 1985 1990 1995 2000 2005
(b)
Figure 7 Changes in grain (cereals and soybean) imports and exports
in Asia (a) and China (b) from 1961 to 2001.
needs, urban consumer needs, and development of grain-
processing facilities in producing-yet-importing markets
(China) and exporting but price-distorting markets (the US).
Recent Chinese restrictions on soybean imports have increased
domestic prices, benefiting farmers in Heilongjiang, the main
soybean-producing province. Limited supplies of soy meal
have increased prices of pork, and soybean oil has also become
more expensive, affecting urban shoppers across the country.
Rising meat consumption in China has led to development of a
soybean feed-processing industry employing 250 000 people.
Restricted imports, for whatever reason, affect their liveli-
hoods. Chinas estimated consumption of soybean is 33 Mt,
which is projected to rise to 50 Mt within 5 years; in the face of
current production of only  17 Mt. Expectations of free trade
under World Trade Organization (WTO) are commonly under-
mined by nontariff barriers. Clearly Chinese imports of soy-
bean from the US should continue and a mutually
beneficial trade will result. The reasons for the restrictions
(safety standards for genetically modified soybeans, and phy-
tosanitary regulations concerning phytophthora) obviously
have a strong policy overtone. However, they may be placed
in context of the widespread demands from developing econ-
omy nations for US farm subsidies and tariffs to be reduced in
the same spirit of free trade.
381
General Discussion
Exports Net imports
Is Biotechnology the Answer?
Biotechnology can provide plant varieties that are more stress-
resistant, responsive to appropriate inputs, resistant to pests
and diseases (removing the necessity for energy-costly treat-
ments), and are suited to specific end uses. Biotechnology
cannot change food production from a water, dirt, and
energy business into an environmentally stress-free self-
sustaining system. Massive increases in food production are
needed in the next several decades to keep pace with present
demographic trends. Hopefully, population growth will slow
and the new plateau in food production will prove to be
sustainable. In some ways, biotechnology is distracting govern-
ments from the true nature of the problem and from their
Years
obligations to act on realistic solutions. Biotechnology has
served poorly if at all in increasing yields relative to energy
Exports Net imports inputs. Yield increases over the 40 ongoing years of the Green
Revolution have been due to development of varieties that can
better sustain such inputs. Limits to population and limits to
demand are keys to global food security. Food supply has
become too politicized for governments to feel secure placing
their fundamental food security in the hands of the most
efficient producers. Perhaps the time has come to develop an
international treaty guaranteeing free and unimpeded trade in
basic foods and medicines, no matter what the prevailing
conditions of political conflict or even war between nations.
The Future
Years Asia is the key to world prosperity, food security, and stability;
or the alternative of massive regional famines. Large increases
in grain prices will not much affect consumers in the industri-
alized west. Like oil-price fluctuations, these will be absorbed
into the economy. Grain raw material prices are a small per-
centage of most consumers monthly food costs in these coun-
tries. Most Asian countries will be able to absorb any such
structural increases in grain prices albeit with some economic
pain during the readjustment. Most of the poorer Asian coun-
tries are more or less self-sufficient in grain, or with increasing
prices will have the motivation and capability to become self-
sufficient. Rapidly industrializing and developing economies
such as China will combine continued substantial domestic
grain production with the capacity to import additional needs
through the strength of their export-focused manufacturing
industries. Perhaps long-term grain price increases will enable
major producers (in North America and Europe) to remove
farm subsidies and move to more effective free-market produc-
tion. An additional 100 Mt, for example, of grain entering
world trade (at far higher prices than today) will be an attrac-
tive prospect to marketing boards and farmers organizations
in the US, Canada, and Australia. But these major exporting
countries have to look carefully at the environmental costs of
meeting such increased demand. The food supply crisis,
although arising from western production constraints and
Asian demands, will have its most serious consequences in
countries, many in Africa, which have limited production,
rapidly rising populations, and few competitive exports.
Many of these are the same countries that suffered most in
382 GRAINS AROUND THE WORLD | Grain Production and Consumption: China and South-East Asia
the oil price crises of the past few decades. With grain more
than with oil, the impact of the crisis will be more immediate
and drastic on the health and survival of the people of these
countries.
Exercises for Revision
Can and should China feed itself?
Was the Green Revolution really a success? Argue the pros
and cons, focusing on the effect on rural populations in
India.
How much grain is required to produce one kg of each
major meat product?
Exercises for Readers to Explore the Topic Further
How can new storage and supply chain technologies reduce
grain losses from farm to consumer?
What are the prospects for a new Green Revolution based
on genetic engineering?
Why is Vietnam a major rice exporter, yet Philippines strug-
gles to feed itself?
See also: Grains Around the World: Grain Production and
Consumption: Cereal Grains in North America; Grain Production and
Consumption: Europe; Grain Production and Consumption in Oceania:
Australia and Pacific Countries; Grain Production and Consumption:
Overview; Grain Production and Consumption: South America.
Further Reading
Brown L (1995) Who Will Feed China?, p. 163. New York: W.W. Norton and Company.
Cassman KG and Pingali PL (1995) Extrapolating trends from long-term experiments to
farmers fields: the case of irrigated rice systems in Asia. In: Barnett V, Payne R, and
Steiner R (eds.) Agricultural Sustainability: Economic, Environmental and Statistical
Considerations, pp. 6384. London: Wiley.
Garnaut R and Ma G (1992) Grain in China, p. 151. Canberra: East Asia Analytical Unit,
Department of Foreign Affairs and Trade, Commonwealth of Australia.
Kaosa-ard MS and Rerkasem B (2000) The Growth and Sustainability of Agriculture in
Asia, p. 303. Oxford: Oxford University Press.
Khush GS (1999) Green Revolution: preparing for the 21st century. Genome
42: 646655.
Khush GS (2001) Green revolution: the way forward. Nature Reviews: Genetics
2: 815822.
Wittwer S, Yu Y, Sun H, and Wang L (1987) Feeding a Billion, p. 462. East Lansing,
Michigan: Michigan State University Press.
Relevant Websites
apps.fao.org http://apps.fao.org The FAO database on agricultural production and
trade statistics is a comprehensive resource and invaluable to researchers working
in this area. It has been extensively used in preparation of this article.
irri.org http://www.irri.org Website of the International Rice Research Institute,
Philippines. Information and links on sustainable rice production in Asia.
worldwatch.org http://www.worldwatch.org Website of Worldwatch Institute, for
independent research for an enviornmentally sustainable and socially just society.
 Grain Production and Consumption: Europe
MG Lindhauer, Max Rubner-Institut (MRI), Detmold, Germany
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Europe is one of the most important cereal producers of the
world.
Within Europe, the European Union is the dominant cereal
producer.
Soft (bread) wheat and maize, and to a lower extent barley,
rye, oats, and triticale, are the most important grain cereals.
In some parts of Europe, rye is the basis for bread making.
Other cereals are rice, sorghum, millet, and buckwheat
(pseudocereal).
The per capita consumption of cereals varies significantly
between European countries.
Cereal consumption and preference for individual cereals
are very much dependent on cultural traditions in diet.
Europe is an important producer of pulses (beans, broad
beans, peas, and lentils) and to a lower extent of soybeans.
Rapeseed and sunflower are important oilseeds.
The majority of grains harvested are used for animal feed.
Learning Objective
To get an overview of the agronomically important grain
species grown in Europe and of their importance in human
consumption and other utilization.
Introduction
In Europe, grain crop production has a long tradition, and
grains of different plant origins are substantial constituents of
the human diet and of animal feed since ancient times.
For the cultivation of grain crops, of cereals especially,
Europe has excellent prerequisites because of its climatic and
pedological conditions and its overall efficient agricultural
systems.
Preference for specific grain crops varies very much among
European countries and larger geographic regions owing to the
wide cultural multiplicity and ethnic diversity in nutritional
habits.
Rye may serve as an example. It is a cereal very specific to
Europe and hardly grown not only anywhere else in the world
but also within Europe; it is projected as a bread-making cereal
and animal feed. It is grown almost exclusively in central,
northern, and northeastern countries of Europe.
Another point of consideration while recording statistical
data is that because the European Union (EU) is politically very
diverse, it dominates all the markets including the agricultural
one. This is the case, although the EU as yet does not include
countries like Russia and other countries of the former Soviet
Union.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00047-4
Consequently, data have been prepared and presented with
respect to this given political situation but concentrating more
on the main players to ensure clarity.
Grain Production
Cereals
In Europe, cereals are by far the most important agricultural
crops (Tables 14) and among these, wheat is the principal
crop (Table 1). Wheat accounts for nearly half of the total
quantity harvested in 2012. Grain maize and barley and, to a
distinctly less degree, oats, rye, and triticale are also of signifi-
cance. Triticale, the result of crossbreeding wheat and rye, is in
demand in those cultivation areas where normally rye is grown
and where farmers are looking for alternatives.
All other cereals are of minor importance. The pseudocereal
buckwheat a plant belonging botanically to the Polygonaceae
family was grown throughout Europe until the early 1900s. It
had mostly been grown in areas with poor soils and served for
the production of grits and for specific traditional dishes such
as pancakes and crepes. It has survived as a kind of specialty
crop, for example, in Northern Germany, Russia, Austria, and
individual Balkan states. In very recent years, there has been a
certain rekindling of interest in buckwheat as a functional food
in modern nutrition systems because of constituents such as
dietary fibers that have been scientifically proved to improve
health and avoid diseases in human (animal) metabolism.
Rice is produced only in those parts of Europe that have a
suitable climate, for example, Spain, southern France, Italy, or
Southern Russia.
On the European continent, the EU, with its 28 member
countries, is the most important cereal producer accounting for
almost far more than half of the production of all the cereals,
wheat, barley, maize, and rice (Table 2). Most of the triticale is
grown in the EU.
Among EU member countries, France is the main cereal
producer, followed by Germany, Poland, the United Kingdom,
Spain, and Italy (Table 3). Variance of data between individual
harvest years is due to specific climatic conditions as, for
example, in 2010 and 2011 where drought periods during
the growth period in major production countries (e.g., France
and Germany) resulted in significantly lower cereal yield. Also,
it has to be taken into consideration that data generation may
differ between individual member states, making comparison
of data from different sources sometimes difficult.
Growing of cereals is a long-standing tradition in the Rus-
sian Federation and in Ukraine, though both countries are yet
to utilize their full agronomic capacity (Table 4).
Also, when studying production data for single cereals of
Russia and Ukraine (Table 4), the role of wheat as the most
preferred cereal is underlined. Barley is in the second position
383
384 GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe

Table 1 Cereala production (Mt, millions of tonnes) in Europe in the Table 3 Total cereal production (1000 tonnes) in EU member
year 2013 countries in selected years

Wheat, totalb 225.486 Member country 2009 2010 2011 2012

Barley 85.838
Rye 14.945 Austria 5748 5144 4818 4876
Oats 14.526 Belgium 3324 3105 2944 3012
Maize 117.454 Bulgaria 6384 7079 7461 6941
Triticale 13.546 Croatia 3442 3007 2828 2729
Sorghum 1.156 Cyprus 57 66 74 74
Millet 0.635 The Czech Republic 7832 6878 8285 6596
Rice, paddy 3.895 Denmark 10 117 8748 8794 9460
Buckwheat 1.333 Estonia 874 678 772 994
Mixed grain 3.673 Finland 4261 2989 3668 3687
Cereals, nesc 0.665 France 69 862 65 555 63 681 68 335
Total cereals 483.138 Germany 49 748 44 039 41 920 45 397
Greece 4609 3869 4416 4069
a
Including the pseudocereal buckwheat. Hungary 13 579 12 256 13 669 10 299
b
Soft durum spelt wheat, respectively. Ireland 2063 2040 2509
c
Including inter alia amaranth and quinoa species. Italy 15 892 16 941 18 226
Source: FAOSTAT: FAO Statistical Division (2014). Latvia 1663 1436 1412 2125
Lithuania 3806 2797 3262 4657
Luxemburg 189 166 150 153
Table 2 Cereala production (Mt, millions of tonnes) in the European Malta
Union (EU-28) in the year 2013 The Netherlands 2089 1888 1685 1775
Poland 29 827 27 228 26 767 28 544
Wheat, totalb 225.486 Portugal 957 849 972 991
Barley 59.844 Romania 14 801 16 651 20 777 12 563
Rye 10.239 Slovenia 533 569 608 570
Oats 8.414 Slovakia 3330 2554 3714 3036
Maize 65.056 Spain 16 914 18 941 21 167 16 412
Triticale 11.430 Sweden 5250 4287 4646 5106
Sorghum 0.689 The United Kingdom 21 618 20 946 21 485 19 515
Millet 0.113 EU-28 298 161 280 380 291 559 27 7218a
Rice, paddy 2.787
Buckwheat 0.289 a
COCERAL (2013).
Mixed grain 3.652 Source: Federal Ministry of Food and Agriculture (ed.) (2013) Statistisches Jahrbuch
Cereals, nesc 0.584 uber Ernahrung, Landwirtschaft und Forsten der Bundesrepublik Deutschland, 57. Jhrg.
Total cereals 306.487 Landwirtschaftsverlag GmbH Munster-Hiltrup.
a
Including the pseudocereal buckwheat.
b
Sum of soft durum spelt wheat, respectively.
c Table 4 Cereal and pseudocereal production in Mt (millions of
Including inter alia amaranth and quinoa species.
tonnes) in the Russian Federation and Ukraine in the year 2012
Source: FAOSTAT: FAO Statistical Division (2014).
The Russian Federation Ukraine

in Russia, and in the Ukraine, it is maize. Rye has a certain Wheat 37.72 15.76
importance in Russia but nearly none in the Ukraine. Maize 8.21 20.96
Yields for cereals as a whole and for individual cereal spe- Rye 2.13 0.01
cies reveal a higher cultivation efficiency within the EU Oats 4.03 0.01
(Table 5) when compared with respective data calculated on Triticale 0.46
the whole-Europe basis (Table 6) or with data from Russia or Barley 13.95 6.94
Ukraine (Table 7). The reasons for the more efficient agricul- Sorghum 0.05 <0.001
Millet 0.33 0.001
tural system in most of the European member countries are
Rice, paddy 1.05 0.002
multifold. Without a doubt, in the majority of the countries, Buckwheat 0.80 0.002
farmers are well educated and experienced and can rely on Cereals, total 68.77 45.67
well-established governmental and private consulting systems.
They have unlimited access to production means like fertil- Source: FAOSTAT: FAO Statistical Division (2014).
izers, pesticides, high-value varieties, and modern technical
equipment.
After revision of the EU common agricultural market policy wheat, maize, and barley. But due to an increasing demand
in 2007 and thus unlike as in former decades, grain interven- with coinciding high prices achievable on the world market,
tion, that is, buying for a fixed threshold price and storing cereal intervention has nearly lost its importance.
surpluses from the market, has practically come to an end. It In large parts of the EU, the climate and soil are very
has been restricted to the cereal species soft wheat, durum favorable for cereal cultivation. In the main cereal production
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe 385

Table 5 Cultivation area (1000 ha) and average yield (tonnes ha1) countries in Western and Central Europe, precipitation is
a
of cereals in the European Union (EU-28) in the year 2013 rather high and equally distributed throughout the year. Tem-
peratures are moderate, being, on average, not too low during
Species Cultivation area Yield
winter for the growth of winter-type cereals (winter wheat,
Wheat 25 683 5.6 winter rye, etc.). These cereals have a prolonged vegetation
Barley 12 339 4.8 period and are thus higher in yield.
Rye 2583 4.0 In recent years, however, there seems to be a tendency
Oats 2645 3.2 towards more frequent unusual weather events like drought
Maize 9685 6.7 and very high temperature in spring and during the main
Triticale 2676 4.3 growth period, respectively, and more frequent heavy thunder-
Sorghum 137 5.0 storms with huge amounts of rainfall that may be taken as
Millet 61 1.9
indications of climate change, the consequences of which for
Rice, paddy 430 6.5
cereal production in quantity and processing quality are hard
Buckwheat 159 1.8
Mixed grain 1201 3.0 to assess at present.
Cereals, nesb 119 4.9 Also, among EU member countries, yields of different
Total cereals 57 721 5.3 cereals are varying significantly. Data for bread wheat yield in
2013 (data not shown) may serve as an example. The average
a
Including the pseudocereal buckwheat. yield ranged from 8.6 tonnes ha1 in Belgium to
b
Including inter alia amaranth and quinoa species. 1.9 tonnes ha1 in Portugal. In countries like Belgium/Lux.,
Source: FAOSTAT: FAO Statistical Division (2014).
Denmark, France, Germany, Ireland, the Netherlands, and
the United Kingdom, yields much higher than the EU average
Table 6 Cultivation area (1000 ha) and average yield (tonnes ha1)
a are standard. To a limited degree, there is also a variation in
of cereals in Europe in the year 2013
yield within these countries. These phenomena are the result of
Species Cultivation area Yield differences in climate, soil conditions, utilization of produc-
tion means, and a question of the choice of varieties among
Wheat 57 582 3.9 which there is a broad range in genetically determined yield-
Barley 24 569 3.5 forming potential.
Rye 4978 3.0
Oats 6143 2.4
Maize 18 874 6.2 Wheat
Triticale 3418 4.0
Sorghum 328 3.5 Additional detailed information on wheat has been provided
Millet 495 1.3 because of its overall predominance in production and con-
Rice, paddy 648 6.0 sumption. Data presented in Tables 1, 2, and 4 refer to wheat
Buckwheat 1265 1.1 as a single entity. These statistical data do not distinguish
Mixed grain 1208 3.0 between different wheat species. As mentioned in the preceding
Cereals, nesb 183 3.6 text, the so-called soft aestivum wheat (botanical name Triticum
Total cereals 119 791 4.0 aestivum) is the most important wheat species in Europe and
a
Including the pseudocereal buckwheat. worldwide. Two types of T. aestivum wheat are cultivated: winter
b
Including inter alia amaranth and quinoa species. wheat and spring wheat, where winter wheat is by far the most
Source: FAOSTAT: FAO Statistical Division (2014). preferred due to its higher yield performance. Spring-type wheat
are chosen when they specifically fit into the crop rotation
Table 7 Area harvested (1000 ha) and average yield (tonnes ha1) of system or in those countries (e.g., Scandinavia) where the cli-
cereals and buckwheat in the Russian Federation and in Ukraine in the mate in winter and the growth period are too uncomfortable for
year 2012 winter-type wheat. In the main wheat-producing countries, the
contribution of spring wheat to the total wheat harvest is dis-
The Russian Federation Ukraine tinctly less than 1%. T. aestivum wheat are covering the wide
range of feed and food applications, whereas the species T.
Area Yield Area Yield
durum, known as durum or hard wheat, is specifically grown
Wheat 21 278 1.77 5630 2.80 for pasta production. (This does not mean that pasta production
Barley 7641 1.83 3293 2.11 would not be possible from soft wheat, too!)
Rye 1423 1.50 0.3 2.27 Durum wheat is more adapted to warmer climatic condi-
Oats 2853 0.14 0.3 2.09 tions. Consequently, it is preferably grown in countries with
Maize 1938 4.24 4372 4.79 Mediterranean climates, namely, in Italy, France, and Spain
Triticale 0.2 2.08 (Table 8). Since a couple of years, however, durum wheat
Sorghum 0.04 1.04 0.06 2.23 varieties more adapted to moderate climates are successfully
Millet 0.03 1.02 0.02 1.03
grown in smaller quantities in the Czech Republic, in
Rice, paddy) 0.02 0.05 0.03 6.19
Buckwheat 1030 0.77 0.03 0.87 Germany, and in Austria. In regions with suitable growth con-
Cereals, total 36 988 1.86 14 412 3.17 ditions within these countries, durum wheat means a produc-
tion alternative for farmers and a local supply of durum mills
Source: FAOSTAT: FAO Statistical Division (2014). with produce.
386 GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe

Table 8 Cultivation area, yield, and production of durum wheat in the Table 9 Production of selected cereals (1000 tonnes) in Europe in
European Union (EU-28) in the year 2013 2012

Member Area Yield Production Wheata Maizeb Barley Ryec

country (1000 ha) (tonne ha1) (1000 tonnes)
Austria 1276 2351 663 219
Austria 12 5.1 63 Belgium 1835 734 364 3
France 335 5.2 1742 Bulgaria 4455 1718 662 22
Germany 9 5.9 53 Croatia 1000 1341 235 2
Greece 535 1.9 1000 Cyprus 23 48
Italy 1240 3.2 4005 The Czech Republic 3519 928 1617 148
Portugal 7 2.1 15 Denmark 4525 75 4059 384
Spain 342 2.6 903 Estonia 485 342 57
Cyprus 6 1.7 10 France 37 921 15 614 11 348 160
The Czech 40 4.8 192 Finland 887 1578 66
Republic Germany 22 409 5515 10 391 3878
Hungary 12 4.5 54 Greece 1569 2010 326 28
Malta 2 4.6 9 Hungary 4011 4742 996 78
Slovakia 20 4.6 92 Italy 7767 8195 960 312
Romania 5 2.0 10 Ireland 708 1152
Bulgaria 27 3.5 95 Latvia 1540 249 124
EU-28 2592 3.2 8243 Lithuania 2999 79 742 157
Luxemburg 79 2 38 6
Source: COCERAL (2013). Malta
The Netherlands 1302 245 203 9
Poland 8608 3996 4180 3163
Portugal 59 849 21 15
Romania 5298 5949 971 19
Slovenia 188 272 85 3
In some parts of France, Switzerland, Germany, and Austria,
Slovakia 1275 1170 471 49
cultivation of spelt wheat (T. spelta) has been conserved Spain 5190 4115 5977 256
through the centuries for specific use in certain foods and for Sweden 2289 11 1705 140
brewing a special beer. In recent times, people have become The United Kingdom 13 261 23 5522 33
more and more conscious of the important role of a healthy EU-28 134 476 60 079 54 901 9332
nutrition, and it is of benefit for health to consume multifold Iceland 15
foods and among them those with functional constituents Norway 235 550 5
(such as dietary fiber/functional food), and in this context, Switzerland 512 149 185 11
emmer (T. dicoccum), small spelt (T. monococcum), and Kamut The Republic of Macedonia 215 119 90 7
Turkey 16 807 4600 7100 371
(T. polonicum) are gaining certain renewed recognition. This is
more so in the case of organic farming and its niche clientele. a
Including spelt wheat.
However, the production rates are negligibly low. In addition, b
Kernel maize including corn cob mix.
only baked spelt wheat goods, mainly bread, are more fre- c
Including maslin.
quently available in specialized bakeries. Source: EUROSTAT (2013); Federal Ministry of Food and Agriculture (ed.) (2013)
Statistisches Jahrbuch uber Ernahrung, Landwirtschaft und Forsten der Bundesrepublik
Deutschland, 57. Jhrg. Landwirtschaftsverlag GmbH Munster-Hiltrup.
Rye
According to the 2012 databases, Germany is the most
Rye is a very important cereal for Europe and more precisely important rye producer, followed by Poland (Table 9) and
for central, east, and northeast European countries, and it is Russia (Table 4). In the Baltic states and in the Nordic
hardly grown anywhere else in the world (Tables 1, 2, and 4). countries, rye and rye products are important constituents of
This has a historical basis. Rye is assumed to have originated the human diet and animal feed, though also in these
from Southwest Asia (Turkey, Iran, and Iraq). From there, it is countries, there is a tendency to lower rye in favor of increased
said to have moved to Central Europe as a weed along with wheat consumption.
wheat and barley. But ancient farmers learned to appreciate this
weed as it turned out to be often more stable and efficient in
Pulses and Oilseeds
yield performing better than the originally sown cereal, espe-
cially on poor sandy soils. Rye has a better drought resistance In comparison with cereals, the production of pulses is of lesser
and efficiency of nutrient uptake from poor substrates. Also, it is importance in Europe (Tables 1012). It is mainly the EU, the
much more tolerant of low temperatures during winter. Russian Federation, Ukraine, and some other European coun-
Consequently, rye was very widely grown in Europe. In tries where the production of specific legumes plays a remark-
some parts, for example, in the United Kingdom, it has been able role. Table 11 provides details of the highest amounts
replaced again by wheat. But in other parts, it remained of being reported for individual pulses and, for some countries,
utmost importance in the human diet, primarily for bread soybeans. Preference for individual species differs very much
making. between countries, depending strongly on traditional diet as
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe 387

far as human consumption is concerned and on recipes of Both plants are of tremendous significance, whereas other
optimized feedstuff mixtures in modern animal production. species like linseed, for example, are specialty crops in specific
At this point, it should be mentioned that there is a strong countries. Basically, for oilseeds, too, it has to be stated that
tendency not only in Europe, namely, in Russia and Ukraine, they are of lesser importance, although they may contribute to
but also in Mediterranean countries to grow soybeans. Even in a large extent to agricultural production as in France, Germany,
countries like Austria or Germany where the climate was con- or the United Kingdom, for instance.
sidered so far to be too unfavorable for this species, attempts
are made to establish soybean production. Making use of
respective progress in breeding the underlying idea is to have Consumption and Utilization
a further feedstuff raw material from home production. The Consumption and utilization of grain seeds in Europe are
majority of the huge amounts of soybeans used, however, will greatly influenced by factors as varied as climate, soil condi-
remain to be imported from overseas. tions, preference for specific crops based on tradition and
Soybean may be used not only as protein source for food ethnic peculiarities, economics, and governmental regulations.
and feedstuff but also as oil production raw material. The These factors are further compounded by the diversity in
statistics, however, reveal rapeseed and sunflower as oilseeds regions and countries found in the European continent at
of importance in European agriculture (Tables 12 and 13). large. As it is impossible to discuss every grain species in detail
or take into consideration the entire range of diversities, an
attempt will be made to focus on major trends and to outline
Table 10 Production of pulses and soybeans (1000 tonnes) in total the main fields.
in Europe, the European Union (EU-28), Russia, and Ukraine in the year Nearly all grain seeds or components of them serve as food
2012 and feed raw materials, and these are by far the most important
fields of utilization. Furthermore, these crops or components
Europe EU-28 Russia Ukraine
derived thereof may be used as basic materials for chemical or
Total pulses 6485 3173 2220 473 technical purposes in the nonfood or feed industry. For exam-
Beans, dry 488 139 7 <1 ple, starch is applied for the synthesis of adhesives.
Chickpeas 89 36 <1
Cowpeas 23 <1
Lentils 91 62 28 <1 Consumption of Cereals
Peas, dry 3420 1272 1160 <1
Pulses, nesa 1186 850 321 <1 Statistically, the EU as a whole is a self-supplier of cereals.
Soybeans 5515 863 1806 2410 Individual countries, however, especially the main grain-
producing ones like France (200%), Germany ( 140%),
a
Including inter alia: hyacinth bean (Dolichos ssp.), jack or sword bean (Canavalia and the United Kingdom (125%), produce much more
spp.), and winged bean (Psophocarpus tetragonolobus). cereal in total than can be consumed by the national popula-
Source: FAOSTAT: FAO Statistical Division (2014). tion. However, there are also countries within the EU like, for

Table 11 Production of pulses and soybeans (1000 tonnes) in selected European countries in 2013

Beans Broad beans Lentils Peas Total pulses Soybeans

Albania 27.0 0.2 30.02 0.10

Austria 13.65 17.60 41.39 82.78
Belgium 0.82 2.20 1.50 4.52
Bulgaria 1.13 1.30 2.00 6.80 0.60
The Czech Republic 1.80 30.70 40.16 13.47
Denmark 14.60 24.60
Finland 10.50 10.50
France 7.55 245.55 22.73 498.94 788.92 110.28
Germany 59.70 129.50 228.80 2.00
Greece 25.00 3.32 3.20 0.75 46.77 4.00
Hungary 0.22 0.20 0.02 43.60 44.37 82.10
Italy 8.53 49.42 1.54 8.60 118.28 472.4
Poland 31.33 0.28 33.23 375.71
Portugal 2.37 20.00 23.07
Serbia 33.79 31.70 109.64 385.21
Spain 10.00 28.10 24.00 177.70 373.30 1.40
Sweden 1.05 40.70 41.75
Switzerland 1.10 12.30 13.59
The United Kingdom 200.00 107.00 737.00
Russia 7.06 7.56 16.63 1350.17 2084.34 1636.00
Ukraine 25.80 4.00 0.70 275.00 371.50 2774.30

Source: FAOSTAT: FAO Statistical Division (2014).

388 GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe

example, Italy (40%) or Spain ( 70%) that lack sufficient of cereals, however, being much dependent on yearly fluctua-
production. This means a compensation of surpluses and tions in total production quantities.
demands, respectively, not only within the EU but also from The majority of the yearly cereal harvest is used for feed-
outside. As a whole, the EU is one of the big players in world- stuff. Germany may serve as an example: Out of c.26 million
wide cereal trade; this is specifically valid for major grain- tonnes of wheat produced per year round, about 7 million
producing member states like France or Germany where there tonnes is milled for human consumption. Food use is domi-
is a strong pressure for export outside the EU or for alternative nated by bread production and the production of other baked
utilization of cereals. The latter aspect, however, is diversely goods like pastries and cakes. Breakfast cereals also contribute
discussed and may steadily lose importance with respect to to the total consumption of cereals, though in comparison
world population growth and therefore increased demand for with bread making, this segment is small, with a permanently
cereals for human consumption. In more recent years, Russia increasing tendency. The significance of grain food use is dem-
and Ukraine have also become significant cereal exporters onstrated in Table 14 as consumption of cereals per capita and
(especially wheat) on the world market, the available amounts year in selected EU member states. Consumption differs a lot
between individual countries, with, for example, Greece having
extraordinarily high consumption rates. In the large majority
Table 12 Total production of pulses (1000 tonnes) and oil crops
of the European countries, the production of bread or compa-
(1000 tonnes) in Europe in the year 2013
rable foodstuff means production from wheat. This is also true
Pulses Oil crops for central, eastern, and northeastern countries, but here in
addition, rye is also used as a raw material for bread making.
Total Europe 6212 27 240 The extent to which rye is used is quite different between
EU-28 3165 15 137 individual countries. For example, in Germany, 850 000
Russia 2084 5294 tonnes of rye is milled for bread production; however, this
Ukraine 372 5948
tendency is slowly decreasing.
Source: FAOSTAT (2012). Bread making does not only mean utilization of pure rye
flour. Most breads offered on the market are made of mixtures
of rye and wheat flour. In relation to wheat, rye has higher
Table 13 Production (1000 tonnes) of rapeseed and sunflower seed proportions of dietary fiber. Actually, this health benefit is used
in the European Union in the year 2012 in arguing in favor of increased rye consumption again. Health
aspects are also the reason of increased interest in the so-called
Rapeseed Sunflower seed
old wheat species as spelt, emmer, or einkorn.
Austria 149 53 With steadily increasing amounts, wheat and maize serve as
Belgium 48 raw materials for starch production (Table 15). The advantage
Bulgaria 271 1388 of these raw materials is based on marketable by-products like
Croatia 26 90 germ for oil extraction or protein for feed (maize) or gluten for
Cyprus 0 food use (and feed) from wheat. The so-called vital gluten from
The Czech Republic 1109 57 wheat starch plants is used in a wide range of feed and food
Denmark 485 0 applications, as a flour improver in the bread industry, for
Estonia 163
example. The main product of the starch industry, starch, is
France 5483 1575
Finland 73 0 mainly used in food applications (Table 16).
Germany 4821 63
Greece 1 160 Table 14 Per capita consumption of cereals (kg year1) in selected
Hungary 401 1302 European countries in the economic year 2011/2012
Italy 25 186
Ireland 0 Cereals, Soft durum
Latvia 304 0 total wheat Other cereals
Lithuania 633 0
Luxemburg 15 0 Austria 90.9 64.4 26.5
Malta 0 Estonia 62.3 38.0 24.3
The Netherlands 7 0 Germany 95.0 69.9 25.0
Poland 1862 Greece 313.0 308.1 4.8
Portugal 10 Hungary 106.6 89.9 16.8
Romania 159 1389 Latvia 112.0 82.9 29.0
Slovenia 17 0 Lithuania 92.2 74.4 17.8
Slovakia 213 197 Luxemburg 60.0 57.3 2.7
Spain 52 0 The Netherlands 109.9 99.3 10.6
Sweden 323 0 Romania 159.2 120.3 38.9
The United Kingdom 2564 0 The United 80.8 76.9 3.9
EU-28 Kingdoma
a
, Data not available. Data of 200910.
Source: Federal Ministry of Food and Agriculture (ed.) (2013) Statistisches Jahrbuch Source: Federal Ministry of Food and Agriculture (ed.) (2013) Statistisches Jahrbuch
uber Ernahrung, Landwirtschaft und Forsten der Bundesrepublik Deutschland, 57. Jhrg. uber Ernahrung, Landwirtschaft und Forsten der Bundesrepublik Deutschland, 57. Jhrg.
Landwirtschaftsverlag GmbH Munster-Hiltrup. Landwirtschaftsverlag GmbH Munster-Hiltrup.
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe 389

Table 15 Total raw material processed, starch production, and the In European tradition, maize as such is of little interest for
respective contribution of specific plant groups in Europe in selected food use, though actual development reveals the so-called
years Mexican food to become a segment of diet, at least in some
European countries.
Year 2001 2005 2008 2010
Minor quantities of grain maize are used for grits produc-
Total raw material (Mt) 21.2 22.6 21.6 22.0 tion, as raw material for snacks, while the great majority of
Potato (%) 41 39 35 32 grain maize is used in feed.
Maize (%) 31 31 33 34 Because of their b-glucan content with health benefits, oats
Wheat (%) 28 30 32 34 are processed to a certain amount to produce special food like
Starch produced (Mt) 9.0 9.6 9.4 9.9 flakes and extruded products. But the majority of the produc-
Potato (%) 20 18 16 14 tion is used for feed. This is also valid for barley, though a
Maize (%) 46 46 47 48
notable quantity of barley is used for beer brewing. Barley is
Wheat (%) 34 36 37 38
also rich in b-glucan, a reason why it should be treated as
Mt, millions of tonnes cereal, the potential of which for food production is
Source: German Starch Industry Association (2012). underestimated.

Consumption of Pulses and Oilseeds

Table 16 Utilization of starch and starch derivatives in Europe in To a limited extent and varying very much between individual
selected years
areas and countries within Europe, pulses are used for human
2001 2005 2008 2010 nutrition (Table 17). Mostly, they are components of feedstuff
mixtures. This is also true for soybeans. Food use is in oil and
Total utilization (Mt) 8.3 9.0 8.8 9.0 protein (tofu) extraction. The quantity outlined for certain
Native starches (%) 23 23 23 25 countries like Germany, for example, does not necessarily indi-
Modified starches (%) 17 20 21 20 cate home products but may represent imports.
Saccharification products (%) 60 57 56 56
Oilseeds are mainly used for oil production and the
Food uses (%) 55 58 60 62
protein-rich residues serve as feedstuff ingredients. The oil,
Nonfood uses (%) 45 42 40 38
especially of rapeseed, is of high nutritional value and there-
Mt, millions of tonnes fore used in human nutrition. However, the bulk is used for
Source: German Starch Industry Association (2012). technical applications, for example, as energy for motor cars.

Table 17 Food supply quantity (1000 tonnes) and consumption per capita (kg) of pulses and soybeans in selected European countries in the year 2011

Beans Peas Pulses (others) Pulses (total) Soybeans

1000 tonnes kg 1000 tonnes kg 1000 tonnes kg 1000 tonnes kg 1000 tonnes kg

Albania 15.00 4.8 0.00 0.0 0.91 0.3 16.00 5.1 0.04 0.0
Austria 1.60 0.2 3.99 0.5 1.27 0.2 7.00 0.8 14.39 1.7
Belgium 8.23 0.8 13.00 1.2 2.06 0.2 23.00 2.1 0.65 0.1
Bulgaria 12.00 1.6 1.00 0.1 3.46 0.5 16.00 2.2 0.31 0.0
The Czech Republic 1.85 0.2 20.00 1.9 6.42 0.6 28.00 2.6 6.30 0.6
Denmark 0.74 0.1 4.00 0.7 1.00 0.2 6.00 1.1 0.35 0.1
Finland 0.12 0.0 6.00 1.1 0.23 0.0 6.00 1.1 0.28 0.1
France 52.70 0.8 30.18 0.5 44.04 0.7 127.00 2.0 3.53 0.1
Germany 8.00 0.1 59.18 0.7 19.00 0.2 86.00 1.0 52.33 0.6
Greece 30.83 2.8 1.53 0.1 20.50 1.8 53.00 4.8 0.16 0.0
Hungary 7.83 0.8 24.39 2.4 5.63 0.3 35.00 3.5 0.05 0.0
Italy 115.07 1.9 45.76 0.8 130.43 2.2 291.00 4.8 0.63 0.0
Poland 36.67 1.0 40.51 1.1 2.99 0.1 81.00 2.1 0.27 0.0
Portugal 23.03 2.2 3.39 0.3 10.36 1.0 36.00 3.4 0.14 0.0
Serbia 45.62 4.8 42.64 4.4 0.59 0.1 90.00 9.4 0.04 0.0
Spain 46.12 1.0 215.89 4.6 141.44 3.0 403.00 8.7 0.69 0.0
Sweden 2.56 0.3 12.00 1.3 2.83 0.3 18.00 1.9 1.23 0.1
Switzerland 2.00 0.3 10.54 1.3 2.28 0.3 15.00 1.9 1.03 0.1
The United Kingdom 10.00 0.2 118.95 1.9 21.65 0.4 151.00 2.4 5.42 0.1
Russia 260.00 1.8 1.14 0.0 261.00 1.8 5.40 0.0
Ukraine 69.82 1.5 1.09 0.0 71.00 1.6 0.61 0.0

Source: FAOSTAT: FAO Statistical Division (2014).

390 GRAINS AROUND THE WORLD | Grain Production and Consumption: Europe
Exercises for Revision
Which are the main cereals produced in Europe as a whole
and in the EU?
What are the main cereal-producing countries in Europe?
What cereal species is a specialty in Europe and hardly
grown elsewhere in the world?
What are the main pulses grown in Europe as a whole and
in the EU?
What are the main oilseeds produced in Europe?
What are the cereal species preferably used for human
consumption in Europe?
What is the significance of soft wheat in bread making in
Europe?
Are there differences in the significance for bread making
between individual European countries?
Are there differences among countries in the preference for
the use of pulses?
Exercises for Readers to Explore the Topic Further
Compare the significance of cereal grains versus pulses in
human nutrition.
Select different countries for their preference in utilization
of different cereal species for bread making.
Identify regions of Europe where rye is the predominant
bread cereal.
See also: Food Grains and the Consumer: Consumer Trends in
Grain Consumption; Genetically Modified Grains and the Consumer;
Grains and Health: Misinformation and Misconceptions; Labelling of
Grain-Based Foods; Food Grains and Well-being: Functional
Foods: Overview; Grains Around the World: Grain Production and
Consumption: Overview; Non-food Products from Grains: Cereal
Grains as Animal Feed; Utilization of the Whole Cereal Plant to
Maximize its Value; Processing of Grains: Evaluation of Wheat-
Grain Quality Attributes; Grain Quality Attributes for Cereals Other than
Wheat; Starch: Uses of Native Starch; The Basics: The Grain Crops:
An Overview; The Cereal Grains: An Overview of the Family of Cereal
Grains Prominent in World Agriculture; Durum Wheat: Overview; Rye
Grain: Its Genetics, Production, and Utilization; Wheat: An Overview of
the Grain That Provides Our Daily Bread; The Legumes and
Pseudocereals: Grain Legumes and Their Dietary Impact: Overview;
Pseudocereals: Overview; The Oilseeds: Oilseeds: Overview;
Soybean: Overview; Wheat Processing: Gluten and Modified Gluten;
Wheat-Based Foods: Cereals: Overview of Uses: Accent on Wheat
Grain.
Further Reading
FAOSTAT: FAO Statistical Division (2014).
Federal Ministry of Food and Agriculture (ed.) (2013) Statistisches Jahrbuch uber
Ernahrung, Landwirtschaft und Forsten der Bundesrepublik Deutschland, 57. Jhrg.
Landwirtschaftsverlag GmbH Munster-Hiltrup.
 Grain Production and Consumption: Cereal Grains in North America
JA Fox and LL Ward, Kansas State University, Manhattan, KS, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
North Americas relatively low population density and
abundant arable land contribute to its position as a major
grain producer and exporter.
The US dominates global maize production, widely uses
GM maize, and in recent years has further increased pro-
duction to meet demand for bioethanol.
Maize is the most important cereal crop in Mexico (70% of
production area), and wheat is in Canada (60% of produc-
tion area).
North America produces only 2% of the worlds rice, but is
still a significant exporter.
Learning Objective
To provide perspective on the role of North America in
global production and trade for the covered grains.
Introduction
According to worldstat.info, the North American continent
(herein defined as comprising the United States, Canada, and
Mexico) has
17% of the worlds arable land and
6.6% of the
worlds population. Given the relative abundance of arable land
(0.52 ha per person vs. world average of 0.2 ha per person), the
continent is a major producer and exporter of several grain crops
including barley, maize, oats, rice, sorghum, and wheat.
This article updates an earlier Encyclopedia of Grain Science
chapter on the North American grain sector (Lukow and
McVetty) with data covering the 12-year period from 2002 to
2013. Like the earlier chapter, which covered the period from
1992 to 2001, we provide an overview of production, consump-
tion, and trade for the major North American grains. Unless
otherwise noted, all data are drawn from the Food and Agricul-
ture Organization (FAO) FAOSTAT database. We begin with a
brief overview of grain production on the continent and then
provide additional detail for each of the major cereal grains.
Figure 1 shows the total hectares of cereal grains (barley,
maize, oats, rice, sorghum, and wheat) harvested annually in
the United States, Canada, and Mexico between the years 2002
and 2013. During the 12-year period between 2002 and 2013,
an average of 82.2 Mha of cereals was harvested in North
America, accounting for almost 13% of the worlds harvested
area (640 Mha). With higher yields, however, North America
accounted for 20% of world cereal production during the same
period. Within North America, the United States, with an
average harvested area of over 57 Mha, accounted for about
70% of the total harvested area, with Canada having 18% and
Mexico 12%.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00048-6
Maize is the most commonly grown crop on the continent,
accounting for almost half of the area in cereals (Figure 2)
followed by wheat with about one-third. Less than 2% of
North American cereal area is in rice production compared to
almost 25% for the world.
Between 2002 and 2013, the area in North American cereal
production increased by about 6.6 Mha (
8.5%). Most of the
increase (5.7 Mha) occurred in the United States, where the
area in maize (corn) increased from 28 Mha to over 35 Mha.
The net increase in US cereal area was partially a result of a
3 Mha reduction in the Conservation Reserve Program (USDA
Farm Service Agency) and reductions in other crops, primarily
hay (down 2.5 Mha) and cotton (down 1.4 Mha) (USDA,
QuickStats). During the same period, the area in oilseeds (soy-
beans, rapeseed, sunflower, and linseed) in North America, the
main alternative to cereal crops, also increased by about 6 Mha
from 36 Mha in 2002 to 42 Mha in 2013. Most of the net
increase in oilseed area occurred in Canada where the area in
rapeseed (canola) production more than doubled from about
3.6 Mha in 2002 to over 8 Mha in 2013.
Figure 3 shows total cereal production in North America
between 2002 and 2013. During that period, production
increased from 361 million tonnes (mt) to 535 mt, an increase
of over 48%. Production increased in all three countries: by 47%
in the United States, by 86% in Canada, and by 15% in Mexico.
On average, the United States accounted for 82% of North
American cereal production over the 12-year period, with
Canada contributing 11% and Mexico 7%.
Maize (Corn)
Table 1 provides data on North American and world maize
production from 2002 to 2013. Maize is the most important
cereal grain in North America accounting on average for 49%
of harvested cereal acres and 71% of cereal tonnage produced.
Maize is also the worlds top cereal crop with over 1 billion
tonnes produced in 2013.
On average, North America produced almost 42% of the
worlds maize between 2002 and 2013, on 25% of the worlds
harvested maize area. Yields in North America averaged
8.2 mt ha1 compared to the overall world average (including
North America) of 4.9 mt ha1. The United States is the worlds
leading producer, accounting for 90% of North American pro-
duction, with Canada contributing 3% and Mexico 7%. US
yields averaged 9.3 mt ha1 over the 12-year period, compared
to 8.6 mt ha1 in Canada and 3.0 mt ha1 in Mexico. Drought
in the US Midwest during the 2012 growing season resulted in
yields declining to 7.7 mt ha1, the lowest average yield during
the 12-year period. Average yields trended upward in each
country during the 12-year period, by 0.03 mt ha1 per year
in the United States, by 0.18 mt ha1 per year in Canada, and
by 0.04 mt ha1 per year in Mexico.
391
392 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America

Harvested area cereals

100 000 000

90 000 000
80 000 000
70 000 000
60 000 000
50 000 000
40 000 000
30 000 000
20 000 000
10 000 000
0
2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013
US Canada Mexico Total N.Am.
Figure 1 Harvested area of barley, maize, oats, rice, sorghum, wheat (hectares).

Cereal area distribution

60%
N. America
49% World
50%

40% 36%
34%

30%
25% 25%

20%

10% 8%
6% 5% 7%
2% 2% 2%
0%
Barley Maize Oat Rice Sorghum Wheat
Figure 2 Cereal area distribution: North America versus world.

Cereal production
600 000 000

500 000 000

400 000 000

300 000 000

200 000 000

100 000 000

0
2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013

US Canada Mexico Total N.Am.

Figure 3 Production of barley, maize, oats, rice, sorghum, and wheat (tonnes).

Since the introduction of genetically modified (GM) crops
in the late 1990s, both the United States and Canada have
experienced widespread adoption of GM maize. In 2013, GM
varieties of maize accounted for 90% of the planted area in the
United States (USDA, ERS) and 98% of the area in Canada
(GMO Compass). GM maize is not grown commercially in
Mexico and its possible introduction is the subject of ongoing
legal challenges (Nature.com).
The area of maize harvested in the United States increased
by over 25% between 2002 and 2013, from 28 Mha to over
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America 393

Table 1 Maize production: Area harvested (000 ha) and production (000 mt)

The United States Canada Mexico North America World

Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 28 057 227 765 1283 8999 7120 19 298 36 460 256 062 137 609 604 872
2003 28 710 256 227 1226 9587 7521 20 701 37 457 286 516 144 701 645 165
2004 29 798 299 874 1072 8837 7688 21 670 38 558 330 381 147 454 728 971
2005 30 399 282 261 1085 9332 6606 19 339 38 089 310 932 148 035 713 682
2006 28 587 267 501 1061 8990 7295 21 893 36 942 298 384 146 741 706 847
2007 35 014 331 175 1369 11 649 7333 23 513 43 716 366 337 158 390 790 115
2008 31 797 307 142 1169 10 592 7354 24 320 40 319 342 054 162 689 830 611
2009 32 169 332 549 1142 9561 6223 20 143 39 534 362 253 158 743 820 203
2010 32 960 316 165 1203 11 715 7148 23 302 41 311 351 181 164 030 851 271
2011 33 990 313 949 1202 10 689 6069 17 635 41 260 342 273 172 257 887 855
2012 35 359 273 820 1418 13 060 6924 22 069 43 701 308 949 178 552 872 792
2013 35 478 353 699 1480 14 194 7096 22 664 44 054 390 557 184 192 1 016 736
Avg. 31 860 296 844 1266 10 600 7031 21 379 40 117 328 823 158 616 789 093

35 Mha, with its share of US cereal area increasing from 52% to
over 60%. Higher yields, in conjunction with the increase in
area, resulted in a 55% boost in production over the same
period. The acreage increase can be attributed largely to
increased demand for ethanol production spurred by the US
renewable fuels mandate introduced in the 2005 Energy Policy
Act (US Environmental Protection Agency) and the attendant
increase in maize price. US maize production is concentrated
in the Midwest corn belt states, with Iowa and Illinois typi-
cally accounting for around one-third of the US crop. The
recent expansion has boosted production in other states such
as Kansas where the area under maize increased from 1 million
to 1.6 million hectares between 2002 and 2013 (USDA, Quick
Stats). Approximately 15% of US corn harvested area is irri-
gated (USDA, Census of Agriculture).
Maize is by far the most important cereal crop in Mexico,
accounting for over 70% of the area under cereals, with pro-
duction occurring throughout the country. The total area in
maize was stable during the period under review, with almost
identical areas harvested in 2002 and 2013. Higher yields
resulted in a 17% increase in production between 2002 and
2013 but productivity gains have been limited by structural
problems. More than 85% of Mexicos 2 million corn growers
have fewer than 5 ha and have limited access to improved
genetics, fertilizer, etc. (USDA, FAS). In Canada, maize is a
relatively minor crop, accounting for an average of only 8.2%
of the area in cereals, with production concentrated in Ontario
and Quebec (Statistics Canada). The area in maize increased
slightly from 1.28 Mha in 2002 to 1.48 Mha in 2013, but
higher yields contributed to an overall increase in production
of about 58%.
Consumption and Trade
Table 2 provides 10-year average balance sheet data on pro-
duction, consumption, and trade for maize in North America
for the period 2002 to 2011. In the balance sheet, domestic
supply is calculated as production imports stock change-
imports. Thus, a negative stock change corresponds to a reduc-
tion in domestic supply. Because total supply accounts for
stock change, the total use number corresponds to total supply.
Using FAO definitions for the various use categories, feed refers
to livestock feed, processing is the amount used for manufacture
of processed food items (e.g., high-fructose corn syrup), food
comprises amounts not detailed otherwise in the balance sheet
that are available for human consumption, while other uses
comprise quantities used for the manufacture of nonfood
products (e.g., ethanol).
In the United States and Canada, maize is grown primarily
for livestock feed and ethanol with relatively small amounts
used for human food consumption. In Mexico, however, white
maize targeted toward human consumption (e.g., tortillas)
accounts for two-thirds of production (USDA, FAS). Over the
10-year period from 2002 to 2011, food use accounted for 63%
of maize produced in Mexico, compared to 6% in Canada and
only 1.3% in the United States. Mexico has traditionally been a
major importer of feed corn for its domestic livestock sector,
particularly the poultry sector, with imports coming primarily
from the United States. Over the 10-year period from 2002 to
2011, average imports of 7.3 mmt were equal to
34% of
average domestic production. In 2002, Mexico imported
5.2 mmt and ranked as the worlds third largest importer
after Japan and South Korea. In 2011, imports had doubled
to 11.2 mmt, with Mexico as the #2 ranked import market after
Japan.
Between 2002 and 2011, Canada was a net maize importer,
but as maize production increased, the country gradually
shifted from being a relatively small net importer to a relatively
small net exporter. In 2002 and 2003, Canadian net imports
amounted to 3.6 mmt and 1.7 mmt, respectively, while in
2011, net imports had fallen to 0.4 mmt. For 2012 and 2013,
Canada was estimated to have net maize exports of between 1
and 1.25 mmt.
In the United States, the growth of the ethanol sector during
the past decade consumed an increasing share of the US maize
crop. Of the US maize crops produced in 2002 and 2003, an
average of 27 mmt (11% of total usage) was used for ethanol
production. Of the 2010 and 2011 crops, the average amount
used for ethanol was 127 mmt, equivalent to 40% of total use
(USDA World Agricultural Outlook Board). The United States
has long been the worlds top-ranked maize exporter and,
despite increasing ethanol production, has maintained the
top export ranking due to increased domestic production.
Throughout the 10-year period, the United States, on average,
394 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America

Table 2 Maize production, trade, and use (mt)

The United States Canada Mexico North America

Production 293 460 800 9 995 000 21 181 430 324 637 230
Imports 566 349 2 562 245 7 283 827 10 412 421
Stock change 2 678 290 (94 500) 280 889 2 864 679
Exports 50 552 700 641 963 308 182 51 502 844
Domestic supply 246 152 739 11 820 778 28 437 950 286 411 467
Use
Feed 138 788 400 9 357 673 10 081 009 158 227 082
Seed 555 280 12 840 396 066 964 185
Waste 371 117 3 208 635 3 579 752
Processing 22 336 530 1 213 531 1 409 676 24 959 737
Food 3 892 730 647 500 13 296 600 17 836 830
Other uses 80 579 890 218 127 45 968 80 843 984
Total use 246 152 830 11 820 788 28 437 953 286 411 570

10-year average (20022011).

Table 3 Wheat production: Area harvested (000 ha) and production (000 mt)

The United States Canada Mexico North America World

Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 18 545 43 704 8711 15 961 635 3236 27 890 62 901 213 817 574 743
2003 21 474 63 803 10 215 23 049 605 2716 32 293 89 568 207 700 560 129
2004 20 222 58 697 9389 24 796 517 2321 30 128 85 814 216 569 632 144
2005 20 277 57 242 9404 25 748 635 3015 30 315 86 005 219 573 626 739
2006 18 940 49 216 9682 25 265 646 3378 29 268 77 860 211 195 602 332
2007 20 639 55 820 8636 20 054 692 3515 29 967 79 390 216 710 612 874
2008 22 541 68 016 10 032 28 611 802 4019 33 374 100 647 222 279 683 207
2009 20 191 60 366 9638 26 848 828 4116 30 658 91 329 224 634 686 720
2010 19 271 60 062 8269 23 167 679 3677 28 218 86 906 216 965 649 325
2011 18 496 54 413 8544 25 261 662 3628 27 702 83 302 220 196 699 389
2012 19 798 61 677 9497 27 205 579 3274 29 874 92 157 217 320 671 497
2013 18 274 57 967 10 442 37 530 634 3357 29 350 98 854 218 461 713 183
Avg. 19 889 57 582 9371 25 291 659 3354 29 920 86 228 217 118 642 690

exported around 17% of domestic maize production. During
the 2002/03 crop marketing year, the United States exported
40.3 mmt compared to total world exports of 78.2 mmt, a
market share of 51.5%. The second ranked exporter was
Argentina with 11.2 mmt. During the 2011/12 crop year, US
exports of 39.2 mmt accounted for a 50.2% share of the
77.8 mmt export market. The second ranked exporter was
Brazil with 24.3 mmt, followed by Argentina with 17.1 mmt.
Wheat
Following maize, wheat is the second most commonly grown
cereal in North America, accounting for about one-third of the
area in cereals. Worldwide, average annual wheat production
between 2002 and 2013 was
640 mmt, with North America
accounting for around 13%.
Table 3 provides data on North American and world wheat
production from 2002 to 2013.
Between 2002 and 2013, the area harvested in North
America remained relatively stable at about 30 Mha, peaking
at 33.3 Mha in 2008 with production peaking at over 100 mmt
in the same year. Since the early 1990s, however, the area in
wheat has dropped significantly with the 29.5 Mha harvested
in 2013 representing a decline of over 25% on the 40.1 Mha
harvested in 1992. Between those years, the US area in wheat
declined by about 7 Mha, and the Canadian area by about
3 Mha.
Wheat is the primary cereal crop in Canada, occupying over
60% of the area in cereals and over 20% of the total arable
land. Most of Canadas wheat production is in the Prairie
Provinces comprising Alberta, Saskatchewan, and Manitoba.
Canada Western Red Spring (CWRS) is the most commonly
produced class accounting for about 60% of the wheat area,
followed by Canada Western Amber Durum (CWAD) with
around 15%. The United States harvested an average of just
under 20 Mha of wheat during the 12-year period, equivalent
to about one-third of the countrys cereal acreage and slightly
more than 10% of the arable land. In 2013, about one-third of
US wheat was hard red winter (HRW), grown primarily in the
Central and Southern Great Plains. Hard red spring (HRS),
grown in the Northern Great Plains, and soft red winter,
grown in the eastern and southeastern states, each accounted
for about a quarter of production. White wheat accounted for
around 12% of production, and durum around 3%. Wheat is a
relatively minor crop in Mexico, occupying only about 7% of
the area in cereals and <3% of arable land, with durum being
the predominant class (USDA, FAS).
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America 395

In 2013, the United States was the worlds fourth largest
producer (after the EU, China, and India) and, with annual
production of around 57 mmt, generally accounts for about
two-thirds of North American production. Of the remainder,
Canada, with an average of about 25 mmt, accounts for about
30%, with only 4% produced in Mexico. A bumper crop in
Western Canada in 2013 saw the Canadian share of North
American production rise to 38%.
North American wheat yields averaged 2.8 mt ha1 over the
12-year period, slightly below the world average of
2.9 mt ha1. Within North America, average yields were high-
est in Mexico at 5.1 mt ha1 compared to 2.9 mt ha1 in the
United States and 2.7 mt ha1 in Canada. Higher yields in
Mexico are due to the fact that much of Mexican wheat pro-
duction is done under irrigation. Average yields trended
upward in each country during the 12-year period, by
0.05 mt ha1 per year in the United States, by 0.1 mt ha1
per year in Canada, and by 0.07 mt ha1 per year in Mexico.
average exports of over 30 mmt representing more than half
of domestic production, the United States is the worlds lead-
ing wheat exporter, accounting in recent years for a 20% share
of the worlds trade.
Most of the domestic wheat supply is for human food
consumption, with only 13% used for animal feed. Canada is
also a major wheat exporter with average exports of 16.7 mmt
representing over 70% of production. Canada is typically one
of the top five wheat exporters with the other major exporters
being the EU, Australia, and Russia. In contrast to the United
States, almost half of Canadas domestic supply is used for
animal feed. Mexico is a net importer of wheat with average
imports exceeding domestic production. Imports doubled
from an average of 1.8 mmt per year in the period from 1991
to 2000 to an average of 3.6 mmt in period from 2002 to 2011.
Approximately two-thirds of Mexicos domestic wheat supply
is for food use.

Sorghum
Consumption and Trade
Table 4 summarizes wheat production, consumption, and Grain sorghum is grown in Mexico and the United States, but
trade in North America for the period 2002 to 2011. With because maturity requires higher temperatures, there is no

Table 4 Wheat production, trade, and use (mt)

The United States Canada Mexico North America

Production 57 134 020 23 875 980 3 362 165 84 372 165

Imports 3 660 880 684 010 3 611 733 7 956 623
Stock change 898 696 (97 472) 0 801 224
Exports 30 429 360 16 732 450 1 117 632 48 279 442
Domestic supply 31 264 250 7 730 067 5 856 267 44 850 584
Use
Feed 4 128 683 3 663 790 78 109 7 870 582
Seed 2 133 622 905 600 72 173 3 111 395
Waste 22 630 487 267 509 897
Processing 0 14 021 14 021
Food 25 000 220 2 796 166 3 781 549 31 577 935
Other uses 1703 382 608 1 437 168 1 821 479
Total use 31 264 229 7 784 814 5 856 265 44 905 308

10-year average (20022011).

Table 5 Sorghum production: Area harvested (000 ha) and production (000 mt)

The United States Canada Mexico North America World

Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 2883 9163 0 0 1744 5206 4627 14 369 41 316 53 496
2003 3156 10 446 0 0 1973 6462 5128 16 908 44 609 59 037
2004 2637 11 523 0 0 1833 7004 4470 18 528 40 832 58 117
2005 2321 9981 0 0 1599 5524 3921 15 505 46 362 59 633
2006 1999 7032 0 0 1600 5519 3600 12 550 43 366 57 772
2007 2748 12 636 0 0 1775 6203 4523 18 839 44 818 62 810
2008 2943 11 998 0 0 1838 6611 4781 18 609 45 382 66 524
2009 2234 9728 0 0 1691 6108 3924 15 836 40 945 56 938
2010 1948 8779 0 0 1768 6940 3716 15 720 41 583 60 056
2011 1590 5447 0 0 1728 6429 3318 11 876 42 312 58 243
2012 2005 6272 0 0 1820 6970 3825 13 242 38 158 57 030
2013 2630 9882 0 0 1689 6308 4319 16 190 42 120 61 385
Avg. 2425 9407 0 0 1755 6274 4179 15 681 42 650 59 253
396 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America
commercial grain sorghum production in Canada (AgAnnex).
Some sorghum is harvested for silage, but the area is small,
typically around 5% of the US area. Table 5 provides sorghum
production data for the United States and Mexico.
Between 2002 and 2013, sorghum occupied about 5% of
North Americas area in cereals, far less than the shares taken by
maize (49%) and wheat (36%). It is, however, the second most
commonly grown cereal crop in Mexico, with 1.7 Mha
accounting for about 18% of harvested cereal acreage. As the
US area in maize has expanded, the area in sorghum has fallen,
and the 2.6 Mha harvested in 2013 represented a 46% decline
on the 4.8 Mha harvested in 1992. North Americas area in
sorghum averaged 4.1 Mha between 2002 and 2013, just under
10% of the world total. With higher yields, averaging
3.9 mt ha1 in the United States and 3.6 mt ha1 in Mexico
compared to a world average of 1.4 mt ha1, the continent
accounted for more than a quarter of world production. Inter-
estingly, US sorghum yields declined from an average of
4.1 mt ha1 in the 10-year period from 1992 to 2001 to an
average of 3.9 mt ha1 between 2002 and 2013. During the
same periods, Mexican average yields rose from 3.2 to
3.6 mt ha1.
In most years, the United States and Mexico are ranked
among the top four sorghum-producing nations along with
Nigeria and India, with each typically producing between 10%
and 15% of the worlds total sorghum crop. In 2013, the
United States was the top-ranked producer with 9.8 mmt repre-
senting 16% of world production, while Mexico ranked third
with 6.3 mmt. In the United States, the largest sorghum-
producing states are Kansas and Texas, which jointly account
for almost 80% of production (USDA, Crop Production). In
Mexico, sorghum is produced throughout the country and
approximately one-third of the planted area is irrigated
(USDA, FAS).
Consumption and Trade
Table 6 summarizes sorghum production, consumption, and
trade in North America for the period 2002 to 2011. With
exports averaging 4.6 mmt per year comprising almost half of
domestic production, the United States is the worlds leading
Table 6 Sorghum production, trade, and use (mt)
The United States
Production 9 673 297
Imports 879
Stock change 89 501
Exports 4 650 196
Domestic supply 5 113 491
Use
Feed 3 777 720
Seed 21 971
Waste
Processing 1 068 700
Food 245 100
Other uses
Total use 5 113 491
10-year average (20022011).
sorghum exporter. Over the 10-year period summarized in
Table 6, US exports accounted for 76% of total world trade,
but its share declined throughout the period as domestic use,
including some for ethanol production, increased and exports
from Argentina, the second ranked exporting country, grew
from < 0.5 mmt in 2002 to over 1.8 mmt in 2011. Most of
the US domestic supply is used for animal feed.
Despite being a major producer, Mexico is also the worlds
leading sorghum importer, accounting in some years for more
than half of world imports. Until 2013, when sorghum was
imported from Argentina, imports were sourced exclusively
from the United States. Mexican sorghum use is for animal
feed, with the poultry industry being the major consumer
(USDA, FAS). Canadas domestic supply, which is also used
exclusively for animal feed, is imported from the United States.
Barley
Similar to wheat, barley has both spring and winter growth
forms, but North American production is predominantly of the
spring form (Lukow and McVetty). North America produces
about 11% of the worlds barley supply, and the crop occupies
about 6% of the North American area in cereals. Table 7 pro-
vides data on barley area harvested and production.
Between 2002 and 2013, the EU was the worlds largest
barley producer, accounting for about 40% of production,
with North America contributing about 11%. Other leading
producers included Russia, Ukraine, and Australia. Canada is
the largest producer in North America, supplying about two-
thirds of the continents barley, equivalent to about 7% of
world production. Canada is typically among the top five
producing countries. After wheat, barley is the second most
commonly grown cereal in Canada, occupying about 20% of
the cereal area. Canadian acreage declined by 20% during the
12-year period described in Table 7, and the 2013 harvested
area of 2.6 Mha was 30% below the 1992 figure of 3.8 Mha.
Barley is a relatively minor crop in both the United States
and Mexico, occupying between 2% and 3% of the cereal area.
In the United States, barley is grown primarily in northern
states, with Idaho, Montana, North Dakota, and Washington
combining to account for almost 80% of the US harvested area.
Canada Mexico North America
6 200 727 15 874 024
5026 2 746 408 2 752 313
0 89 501
114 65 4 650 375
4912 8 947 068 14 065 471
4912 8 739 151 12 521 783
28 974 50 945
178 943 178 943
1 068 700
245 100
0
4912 8 947 068 14 065 470
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America 397

Table 7 Barley production: Area harvested (000 ha) and production (000 mt)

The United States Canada Mexico North America World

Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 1669 4940 3342 7468 282 737 5293 13 145 55 270 136 724
2003 1913 6059 4397 12 164 365 1085 6675 19 308 57 736 142 565
2004 1627 6091 3841 12 557 326 932 5794 19 579 57 567 153 905
2005 1323 4613 3634 11 678 306 761 5263 17 052 55 357 138 680
2006 1194 3923 3223 9573 315 869 4733 14 365 56 386 139 522
2007 1417 4575 3998 10 984 286 653 5701 16 211 55 860 134 175
2008 1529 5230 3502 11 781 311 781 5342 17 792 55 533 154 788
2009 1260 4949 2918 9517 239 519 4416 14 985 53 956 151 782
2010 998 3925 2387 7605 268 672 3652 12 203 47 368 123 683
2011 906 3392 2365 7756 218 487 3489 11 635 48 488 132 951
2012 1313 4796 2060 8012 328 1032 3701 13 840 49 573 133 507
2013 1214 4683 2652 10 237 297 594 4163 15 514 49 781 144 755
Avg. 1364 4765 3193 9944 295 760 4852 15 469 53 573 140 586

Table 8 Barley production, trade, and use (mt)

The United States Canada Mexico North America

Production 4 769 556 10 108 280 749 564 15 627 400

Imports 762 430 125 464 365 608 1 253 503
Stock change (81 203) (124 100) 1886 (203 417)
Exports 886 840 2 280 319 8442 3 175 600
Domestic supply 4 563 943 7 829 327 1 108 616 13 501 886
Use
Feed 1 282 117 6 963 198 331 106 8 576 421
Seed 138 151 328 400 16 496 483 047
Waste 1530 25 319 26 849
Processing 2 982 160 529 539 735 115 4 246 814
Food 161 514 6659 580 168 754
Other uses 0 0 0
Total use 4 563 941 7 829 326 1 108 616 13 501 884

10-year average (20022011).

As in Canada, the US area in barley has also declined from
2.9 Mha in 1992, to 1.7 Mha in 2002, and down to 1.3 Mha in
2013, a reduction of over 55% in 22 years. The decline can
largely be attributed to the expansion of corn production. In
Mexico, the area in barley has been stable, averaging
295 000 ha. Average yields are highest in the United States, at
3.5 mt ha1, and lowest in Mexico at 2.5 mt ha1, while Cana-
dian yields averaged 3.2 mt ha1 over the 12-year period. Aver-
age yields increased slightly in the United States and Canada,
by 0.06 and 0.1 mt ha1 per year, respectively, between 2002
and 2013, but showed no growth in Mexico. Overall produc-
tion of barley in North America is trending downward, how-
ever. In 2012 and 2013, average production was 14.7 mmt
compared to an average of 21.8 mmt in 1992 and 1993.
food and seed. Usage for animal feed has declined in step with
reduced production, while the amounts used for processing
have been fairly steady. North America is a net exporter of
barley, with exports accounting for around 12% of average
annual production.
Canada, with exports averaging about 2.2 mmt, is typically
one of the top five exporting countries. Of its remaining
domestic supply, almost 90% is used for animal feed. In the
United States, however, only 30% of the available supply goes
to animal feed, with the majority of the crop going to the
malting sector. In 2011, for example, malting varieties
accounted for 75% of the acreage planted in Idaho (USDA,
NASS, Idaho Barley Varieties) and over 90% of the acreage in
North Dakota (North Dakota Barley Council).

Consumption and Trade

Table 8 summarizes production, consumption, and trade in
barley in North America between 2002 and 2011. During that
period, almost two-thirds of the domestic supply was used for
animal feed, with another third used in processing mostly as
malt for beer-brewing industry, with smaller amounts used for
Oats
Oats, like barley, are minor cereals in North America and, with
an average of 1.9 Mha harvested per year, accounts for < 3% of
cereal acreage. The distribution of oats in North America is very
similar to that of barley, with Canada accounting for 65% of
398 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America

Table 9 Oat production: Area harvested (000 ha) and production (000 mt)

The United States Canada Mexico North America World

Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 833 1684 1379 2911 42 64 2254 4658 12 436 25 424
2003 898 2096 1415 3377 46 94 2360 5567 12 128 26 541
2004 723 1679 1234 3467 67 103 2024 5249 11 663 26 036
2005 738 1667 1271 3283 77 127 2086 5077 11 250 23 697
2006 633 1357 1537 3852 76 152 2246 5362 11 661 22 612
2007 609 1313 1816 4696 72 125 2497 6134 11 990 25 806
2008 567 1294 1448 4273 100 148 2115 5714 11 279 25 863
2009 558 1351 980 2906 72 130 1610 4388 10 195 23 345
2010 511 1178 906 2480 67 111 1484 3769 9096 19 723
2011 380 779 1030 2997 52 51 1461 3826 9638 22 316
2012 423 987 985 2812 47 84 1455 3884 9580 21 312
2013 417 1016 1107 3888 49 91 1574 4995 9759 23 821
Avg. 607 1367 1259 3412 64 107 1930 4885 10 890 23 875

Table 10 Oat production, trade, and use (mt)

The United States Canada Mexico North America

Production 1 439 809 3 424 110 110 464 4 974 383

Imports 1 980 072 35 045 144 953 2 160 070
Stock change (234 800) (47 726) 0 (282 526)
Exports 99 493 1 754 758 390 1 854 641
Domestic supply 3 085 586 1 656 673 255 027 4 997 286
Use
Feed 1 774 544 1 491 643 190 960 3 457 147
Seed 138 004 147 220 3782 289 006
Waste 900 1970 2870
Processing 0
Food 1 173 038 16 909 58 315 1 248 262
Other uses
Total use 3 085 586 1 656 672 255 027 4 997 285

10-year average (20022011).

the area, the United States with 31%, and Mexico < 4%. Table 9
provides data on oat production between 2002 and 2013.
Worldwide, oats accounted for only 1.7% of the area in cereals,
and between 2002 and 2013, the harvested area declined by
more than 20%. North America accounts for about 20% of the
worlds production.
The EU and Russia are the leading oat producers, account-
ing for roughly 35% and 20% of production, respectively.
Canada is the third ranked producer with around 15% of
world production (Agriculture and Agri-Food Canada). In
Canada, oats occupy about 8% of the area in cereals, a share
similar to that of maize. In the United States and Mexico, its
share is much lower at 1.1% and 0.6%, respectively.
Mirroring the trend in world acreage, between 2002 and
2013, the area in oats declined in both the United States and
Canada. The decline has been sharpest in the United States
where the 416 Tha harvested in 2013 represented half the area
harvested in 2002 and less than a quarter of the 1.8 Mha
harvested in 1992. In the United States, oats are grown primar-
ily in the North Central states, with Minnesota, North and
South Dakota, Wisconsin, and Iowa jointly accounting for
more than half of the US harvested area. In Canada, most of
the production is in the Prairie Provinces (Alberta,
Saskatchewan, and Manitoba). Yields in the United States aver-
aged 2.3 mt ha1 between 2002 and 2013, compared to
2.7 mt ha1 in Canada and 1.7 mt ha1 in Mexico. Yields in
the United States and Canada evidenced a slight upward trend
over the period, but as was the case with barley, there was no
similar upward trend in Mexican yields. The area in oats in
Mexico ranged from 42 000 ha in 2002 to 100 000 ha in 2008,
with most production in the northwest of the country.
Consumption and Trade
Table 10 summarizes production, consumption, and trade in
oats in North America between 2002 and 2011. As with barley,
oats are used primarily for animal feed. Of the average domes-
tic supply of
5 mmt, around 3.5 mmt or 70% was used for
animal feed, with most of the remainder used for human
consumption. The amount of oats used for human food has
remained relatively constant averaging 1.3 mmt per year in
the period between 1992 and 2001 and 1.25 mmt between
2002 and 2011. As production declined, the amount used for
animal feed also declined, averaging 3.5 mmt per year between
2002 and 2011, compared to 4.9 mmt between 1992
and 2001.
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America 399

Trade in oats is fairly limited with exports from Canada to
the United States accounting for between 55% and 65% of
annual world trade, making Canada the worlds largest
exporter and the United States the largest importer. Canadas
exports averaged 1.7 mmt, equivalent to half of domestic pro-
duction, while US imports exceeded its own domestic produc-
tion. Mexico also imports more oats than it produces
domestically with imports coming primarily from Canada.
Rice
Data on North American and world rice production (paddy
equivalents) from 2002 to 2013 are provided in Table 11. The
United States accounts for over 95% of North American rice
acreage and production. Mexico harvests about 50 000 ha
annually accounting for < 0.5% of its cereal area. Because the
climate is unsuitable, no rice is grown in Canada.
Combined, the United States and Mexico account for < 2%
of world production. Global production is concentrated in
Asia, with China and India combining for over half the world
total. Another 20% of production comes from Thailand,
Vietnam, Indonesia, and the Philippines.
The US area in rice declined by over 20% between 2002 and
2013, from 1.3 Mha to just under 1.0 Mha. An average yield
increase of 0.08 mt ha1 per year tempered the decline in
production to about 10% over the 12-year period. The leading
rice-growing state is Arkansas with about 44% of the total US
area, followed by California with 22%. Rice is also grown in
Louisiana, Mississippi, Missouri, and Texas. Rice is classified as
either long grain, medium grain, or short grain depending on
the shape and length of the grain. US production is predomi-
nantly long grain, which is grown mostly in Arkansas and the
other southern states and accounts for over 70% of US produc-
tion. Medium and short grain types are grown almost exclu-
sively in California, although Arkansas does have some
medium grain production.
Consumption and Trade
Table 12 provides data on rice production, consumption, and
trade (in paddy equivalent tonnes, not milled) in North America
between 2002 and 2011. Although the United States is relatively
small in terms of global production, it is a major exporter. On
average, over the 10-year period considered in Table 12, the
United States exported over half of its domestic production and,

Table 11 Rice production: Area harvested (000 ha) and production (000 mt)

The United States Canada Mexico North America World

Year Area Prod. Area Prod. Area Prod. Area Prod. Area Prod.
2002 1298 9569 0 0 50 227 1348 9796 147 628 571 388
2003 1213 9067 0 0 60 273 1273 9340 148 509 587 031
2004 1346 10 540 0 0 62 279 1408 10 818 150 554 607 986
2005 1361 10 108 0 0 57 291 1419 10 399 154 989 634 444
2006 1142 8826 0 0 70 337 1212 9163 155 581 641 207
2007 1112 8999 0 0 71 295 1183 9293 155 040 656 978
2008 1204 9241 0 0 50 224 1255 9466 159 993 688 415
2009 1256 9972 0 0 54 263 1310 10 235 158 130 686 958
2010 1463 11 027 0 0 42 217 1505 11 244 161 189 701 999
2011 1059 8389 0 0 34 173 1093 8562 162 800 726 122
2012 1084 9051 0 0 32 179 1116 9230 162 317 738 188
2013 1000 8613 0 0 33 180 1033 8793 164 722 745 710
Avg. 1211 9450 0 0 51 245 1263 9695 156 788 665 535

Table 12 Rice production, trade, and use (mt)

The United States Canada Mexico North America

Production 9 573 761 257 963 9 831 724

Imports 1 164 158 611 334 843 477 2 618 969
Stock change (122 019) (5) (794) (122 817)
Exports 5 482 877 147 972 43 982 5 674 831
Domestic supply 5 133 021 463 357 1 056 664 6 653 042
Use
Feed 0
Seed 173 115 3451 176 565
Waste 938 409 13 457 48 726 1 000 591
Processing 694 156 74 417 768 572
Food 3 129 554 420 033 927 595 4 477 181
Other uses 213 814 29 868 2477 246 159
Total use 5 149 047 463 358 1 056 665 6 669 069

10-year average (20022011).

400 GRAINS AROUND THE WORLD | Grain Production and Consumption: Cereal Grains in North America
with an 11% share of world exports, was the fourth largest
exporter following Thailand, Vietnam, and India and ranking
marginally ahead of Pakistan. Mexico is the largest export market
for US rice with other important markets including Japan, Turkey,
and a number of Central American and Caribbean countries.
The United States also imports rice, but imports are mostly
aromatic varieties that are not grown domestically, such as
jasmine from Thailand and basmati from India and Pakistan
(USDA, ERS, Rice). Canadas domestic supply is all imported
with about half coming from the United States. Mexico is also
reliant on imports, which account for close to 80% of its
supply. Those imports come primarily from the United States,
but Mexico also imports from Uruguay and Pakistan (USDA,
FAS, Mexico Grain and Feed Annual). In North America, rice is
used predominantly for human food with some use in food
manufacturing (rice starch, beer, or alcoholic beverages).
Exercises for Revision
Use the data in the tables to construct pie charts describing
the 2013 allocation of cereal hectares among the six grains
for each of the three North American countries.
Again, using the data provided, create line graphs showing
the trend for the North American share of world production
for each of the six grains between 2002 and 2013. For which
of the grains is there a noticeable upward or downward
trend in the North American share of global output?
If the decline in US oat production continues at the same
rate as occurred between 2002 and 2013, in what year will
oat production cease in the United States? Within Europe,
what are the leading oat-producing countries?
Compare the trends in world average yields for oats and
barley.
Compare the trends in US, Mexican, and world sorghum
yields between 2002 and 2013. Are there any noticeable
differences? (Hint for a cleaner comparison, first subtract
US and Mexican area and production from the world num-
bers before calculating the average yield for the rest of the
world).
Estimate the trend growth in US maize yields (with and
without the drought year of 2012). What is the predicted
maize yield for 2020 using both models?
Compare the trends in US and Canadian average yields for
maize and wheat.
Exercises for Readers to Explore the Topic Further
Investigate the importance/share of organic production for
each of the major grains. What are the recent trends in
production of organic corn and soybeans in the North
American countries? Is North America a net imported or
exported of organic grains?
Investigate the prospects for rice production, particularly
wild rice, in Canada. Can you find data on wild rice pro-
duction? How significant is production of wild rice in rela-
tion to Canadian rice imports?
Use the FAOSTAT database to obtain data on US, Canadian,
and world area (hectares harvested) and production (tonnes)
of maize for as many years as possible. Subtract the US and
Canadian data from the world data to facilitate comparison
between the United States, Canada, and the rest of the world
(RoW). Divide production by area harvested to obtain aver-
age yield, and plot the average yield over time for the United
States, Canada, and the RoW. Comment on the trends.
Further Reading
Lukow OM and McVetty PBE. Grain production and consumption: Cereal grains in
North America. Encyclopedia of Food Grains, 1st edn.
United States Department of Agriculture (USDA), Foreign Agricultural Service (FAS).
(2013) Global Agricultural Information Network (GAIN) Report Number MX3024.
Grain and Feed Annual, Mexico. http://gain.fas.usda.gov/Recent%20GAIN%
20Publications/Grain%20and%20Feed%20Annual_Mexico%20City_Mexico_3-
15-2013.pdf.
Relevant Websites
http://www.agannex.com/research/sorghum-in-canada Ag Annex Sorghum in
Canada.
http://www.agr.gc.ca/eng/industry-markets-and-trade/statistics-and-market-
information/by-product-sector/crops/crops-market-information-canadian-industry/
market-outlook-report/oats-situation-and-outlook-august-2010/?
id1378843401619 Agriculture and Agri-Food Canada. Oats: Situation and
Outlook.
http://www.agcensus.usda.gov/Publications/2012/Online_Resources/Ag_Atlas_Maps/
Crops_and_Plants/ United States Department of Agriculture (USDA), Census of
Agriculture.
http://barley.idaho.gov/pdf/quality_reports/2011varieties.pdf United States
Department of Agriculture (USDA), National Agricultural Statistics Service (NASS).
Idaho Barley Varieties.
http://www.epa.gov/otaq/fuels/renewablefuels/ United States Environmental
Protection Agency, Renewable Fuel Standard.
http://www.ers.usda.gov/data-products/adoption-of-genetically-engineered-crops-in-
the-us.aspx United States Department of Agriculture (USDA), Economic Research
Service (ERS).
http://www.ers.usda.gov/topics/crops/rice/trade.aspx Adoption of Genetically
Engineered Crops in the U.S.
http://www.fsa.usda.gov/FSA/webapp?areahome&subjectcopr&topicrns-css
United States Department of Agriculture (USDA), Economic Research Service (ERS).
Rice.
http://faostat3.fao.org/download/Q/*/E FAOSTAT, Food and Agriculture Organization
of the United Nations.
http://gain.fas.usda.gov/Recent%20GAIN%20Publications/Grain%20and%20Feed%
20Annual_Mexico%20City_Mexico_3-15-2013.pdf United States Department of
Agriculture (USDA) Farm Service Agency, Conservation Programs.
http://www.gmo-compass.org/eng/agri_biotechnology/gmo_planting/ GMO
Compass.
http://www.nature.com/news/gm-maize-splits-mexico-1.15493 Nature.com GM
maize splits Mexico.
http://www.ndbarley.net/barleystatistics/ North Dakota Barley Council.
http://www.statcan.gc.ca/pub/96-325-x/2014001/article/11913-eng.htm Statistics
Canada. Corn: Canadas third most valuable crop.
http://quickstats.nass.usda.gov/ United States Department of Agriculture (USDA),
National Agricultural Statistics Service (NASS), Quick Stats.
http://usda.mannlib.cornell.edu/MannUsda/viewDocumentInfo.do?documentID1000.
United States Department of Agriculture (USDA) National Agricultural Statistics
Service (NASS), Acreage
http://usda.mannlib.cornell.edu/MannUsda/viewDocumentInfo.do?documentID1046
United States Department of Agriculture (USDA), National Agricultural Statistics
Service (NASS), Crop Production.
http://usda.mannlib.cornell.edu/MannUsda/viewDocumentInfo.do?documentID1194
United States Department of Agriculture (USDA), World Agricultural Outlook
Board. World Agricultural Supply and Demand Estimates, Various Issues.
http://en.worldstat.info/ Worldstat.
 Grain Production and Consumption: Oilseeds in North America
PBE McVetty, University of Manitoba, Winnipeg, MB, Canada
OM Lukow, Agriculture and Agri-Food Canada, Winnipeg, MB, Canada
LM Hall, University of Alberta, Edmonton, AB, Canada
I Rajcan, University of Guelph, Guelph, ON, Canada
H Rahman, University of Alberta, Edmonton, AB, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The major oilseeds grown in North America are canola,
soybean, linseed/flax, and sunflower.
Whereas sunflower is native to North America, the other
species have been introduced.
Statistics are provided for disposition and consumption of
these oilseeds as 10-year means, as well as historical and
recent annual figures for acreage and production for Can-
ada, the United States, and Mexico.
Canola (Brassica napus) has increased in area because it is
adapted to cool Canadas climate and has been bred to have
low levels of erucic acid in seed oil and glucosinolates in
seed meal.
Canada is the major producer of canola with much of it
being exported.
The United States is the major producer of soybeans, with
much of it being used domestically.
Linseed and sunflower are relatively minor crops in North
America.
Learning Objectives
To achieve understanding of the origins and production of
the major oilseed species in Canada, the United States, and
Mexico.
To understand the uses of these species and the extent of
their involvement in world trade.
Introduction
North America is the major production area for oilseeds and is
a primary or secondary producer in the world of many of the
oilseeds reviewed in this article. Major oilseeds grown in this
continent include canola, soybean, linseed/flax, and sun-
flower. These oilseeds are grown, to greater or lesser extent, in
three countries comprising this continent, Canada, the United
States, and Mexico.
Several of the major oilseeds, such as canola, soybean, and
linseed/flax, currently grown in North America were introduced
in the last few centuries, whereas sunflower is native to this
continent. All the current major oilseeds in this continent have
grown from small beginnings to their major field crop status
today based on successful research and development in cultivar
development, production practices, market development, and
new use development. This article will review the area used to
produce each of these major oilseeds and their production in
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00052-8
each North American countries and on the continent since
1960s. The last 10-year mean disposition and consumption of
each of these major oilseeds in each of the countries and on the
continent will also be reviewed. All comments regarding crop
disposition and consumption refer to averages of the 2000s,
unless specifically stated otherwise. Area and production data
for all crops relate to crops harvested for dry grain only. For
disposition and consumption of commodities, the definitions
of the listed elements are taken primarily from the Food and
Agriculture Organization (FAO) of the United Nations. Values
given for production relate to the total domestic production.
Imports cover all movements into the country of the commod-
ity. Stock variation is the changes in stocks at all levels between
the production and the retail levels. A negative sign indicates
increases in stocks (add to stock) and no sign means net
decreases (from stock). Exports cover all movements by the
crop commodity out of the country. Domestic supply is defined
as (production imports)  (exports changes in stocks). Feed
is the amount of the commodity in question fed to livestock,
whether domestically produced or imported. Seed comprises
the amount of commodity used for reproduction. Waste
includes the amount of crop commodity lost in processing,
storage, and transportation. Processing is the amount of com-
modity used for the manufacture of processed products that
cannot be converted back to their originating primary commod-
ities. Food available for human consumption comprises the
amounts of the commodity not detailed otherwise in the bal-
ance sheet. Other uses comprise quantities of the commodity
used for manufacture of nonfood products.
Canola (Double-Low Rapeseed)
Introduction
Rapeseed includes the Brassica oilseed crop species Brassica
napus and Brassica rapa, while Brassica juncea is commonly
called mustard. These oilseed crops collectively contribute
about 15% of the worlds total supply of vegetable oils and
became the third-leading source of oil in the world after soy-
bean and palm. Brassica oilseeds contain about 4046% oil (in
some cases up to 50%) and the seed meal remaining after oil
extraction contains about 40% protein. Worldwide, this meal
is the second most important (after soybean) economical pro-
tein source used in animal feed. Brassica oilseed crops are
primarily grown to produce edible oil and meal for animal
feed; however, small quantities are used directly for feed and
planting seed.
Because of the ability of the Brassica seeds to germinate and
grow at low temperatures, these oilseed crops are among the
401
402 GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America

few oil crops that can be grown in the temperate regions of the Table 1 Area harvested (ha) for major oilseeds grown in Canada
world. Among these, B. napus is predominantly grown in North
American countries; however, B. rapa and B. juncea can also be Year Canola Soybean Linseed Sunflower
found on limited areas (<1%). B. rapa is more suited to short 196367 473 395 104 650 721 569 23 253
growing season areas, while B. juncea is more suitable to the 196872 1 269 886 139 534 830 993 49 768
hotter and drier regions. B. napus exists in both spring and 197377 1 311 010 178 307 532 374 34 845
winter growth habit forms, but it is the spring form that pre- 197882 2 296 424 296 980 621 610 116 520
dominates in North American production; winter form is 198387 2 695 840 403 880 647 360 46 660
grown on very limited areas in the United States. This oilseed 198892 3 060 920 555 400 508 800 58 360
crop was first introduced in Canada and grown at a commer- 199397 4 692 620 855 900 679 520 58 060
cial scale during the Second World War. 19982002 4 653 100 1 027 500 690 180 75 580
2003 4 689 200 1 048 600 728 400 105 600
Traditional rapeseed and mustard seeds contain a high level
2004 4 867 000 1 173 500 518 000 55 400
of erucic fatty acid in oil (>40% of the total fatty acids) and a
2005 5 175 400 1 165 200 732 600 70 800
high level of glucosinolates in seed meal (>100 mmol g1). The 2006 5 238 200 1 201 200 785 200 76 900
erucic acid is undesired in edible oil, while the seed meal 2007 6 327 600 1 171 500 524 000 78 900
containing a high level of glucosinolate is undesirable for use 2008 6 494 400 1 195 400 625 200 68 800
of this meal as protein source in animal feed. The oil and meal 2009 6 513 300 1 383 300 623 300 63 500
quality of these oilseed crops has been dramatically improved 2010 6 848 300 1 476 800 353 300 51 400
(virtually eliminated erucic acid in oil and reduced glucosino- 2011 7 471 300 1 542 400 273 200 13 800
late content to <30 mmol g1 seed meal) in 1960s1970s 2012 8 379 900 1 668 400 360 000 40 000
through research, and the name canola for this quality- 200312 6 200 460 1 302 630 552 320 62 510
improved-type Brassica oilseed crops has been adopted in Italicized 5- or 10-year means.
most parts of the world. The term double-low rapeseed or Source: Food and Agriculture Organization of the United Nations (FAO).
rapeseed or oilseed rape is still being used for this quality-
improved type in some countries as well as in some articles.
Most of the world production of Brassica oilseeds including the Table 2 Area harvested (ha) for major oilseeds grown in the
production acreages in North American countries has trans- United States
ferred to canola quality type; therefore, the word canola will
be used in this article. Year Canola Soybean Linseed Sunflower seed
Currently, most of the canola grown in North American
196367 0 13 775 857 1 078 310 37 669
countries is hybrid cultivars. About half of the canola crop is
196872 0 17 268 909 827 732 146 238
Roundup (glyphosate) herbicide-tolerant and half Liberty
197377 0 21 679 194 571 738 472 853
(glufosinate) herbicide-tolerant; Clearfield (imidazolinone) 197882 0 27 309 878 286 500 1 653 180
herbicide-tolerant crops cover a small acreage (<5%). 198387 3060 24 741 123 230 500 1 078 720
198892 38 960 23 432 403 93 014 831 596
199397 163 604 25 261 600 60 464 1 183 810
Area Harvested
19982002 515 835 29 199 730 202 993 1 157 571
Canada has harvested canola on an average from about 6.2 2003 432 695 29 330 310 237 958 889 104
million hectares (Mha) (14.4% of Canadian arable land) dur- 2004 338 240 29 930 060 206 797 692 425
ing the period of 200312 (Table 1). Since 1980s, canola 2005 451 634 28 834 570 386 480 1 056 240
2006 413 593 30 190 680 310 397 716 300
production area in Canada has increased more than twofold
2007 468 070 25 959 245 141 237 814 236
with a sharp increase in the recent years (>8 Mha in 2012)
2008 400 160 30 222 654 137 595 969 640
(Table 1). The vast majority of canola production area in 2009 329 780 30 906 980 127 070 790 560
Canada is in the Prairie Provinces Alberta, Saskatchewan, and 2010 579 880 31 003 300 169 160 758 310
Manitoba, while small production areas can be found in 2011 422 620 29 856 410 70 010 589 960
Ontario and British Columbia. 2012 700 560 30 798 530 135 980 745 030
The United States started to grow canola from the late 200312 453 723 29 703 274 192 268 802 181
1980s. This country on an average has harvested canola from
Italicized 5- or 10-year means.
about 453 725 ha (0.3% of American arable land) annually
Source: Food and Agriculture Organization of the United Nations (FAO).
during the period of 200312 (Table 2); this is more than a
twofold increase since the 1990s. The vast majority of canola
production area in the United States is the northern state of land) annually in the last 10 years (Table 3). The area of canola
North Dakota where spring canola is grown. Small production harvested in Mexico has varied widely during the years. Canola
areas can also be found in Minnesota, Montana, Idaho, Ore- production area in Mexico has reduced to less than half in the
gon, Washington, and Oklahoma. Recently, winter canola has recent years when compared with 1970s (Table 3). This varia-
captured about 15% of the canola production area in the U- tion in harvested area is typical for any small-volume crop. The
nited States, primarily grown in Oklahoma and small areas in vast majority of canola production area in Mexico is in the
the Pacific Northwest region. central area of the country.
Mexico is a very small canola producer, harvesting on an For the entire North American continent, canola was har-
average from only about 2000 ha (0.01% of Mexican arable vested from an average 6.7 Mha (2.9% of North American
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America 403

Table 3 Area harvested (ha) for major oilseeds grown in Mexico Table 5 Production (tonnes) for major oilseeds grown in Canada

Year Canola Soybean Linseed Sunflower Year Canola Soybean Linseed Sunflower

196367 5800 41 925 17 623 0 196367 429 557 202 003 518 155 15 310
196872 6400 151 696 17 340 13 415 196872 1 257 833 278 264 689 302 41 111
197377 5800 288 624 10 262 3354 197377 1 403 862 374 759 443 500 36 893
197882 3600 297 419 4795 9905 197882 2 697 258 663 476 609 519 142 980
198387 807 421 432 683 9545 198387 3 430 320 978 700 757 940 57 440
198892 177 315 880 85 4098 198892 3 758 120 1 309 720 546 360 85 100
199397 1025 166 454 27 1295 199397 6 120 800 2 260 540 887 780 76 340
19982002 3200 75 084 367 470 19982002 6 636 900 2 438 280 820 740 122 840
2003 1000 67 880 1350 124 2003 6 771 200 2 273 300 754 400 142 300
2004 1000 88 840 2900 90 2004 7 673 600 3 043 900 516 900 52 200
2005 4000 96 266 1400 22 2005 9 483 300 3 155 600 990 600 84 400
2006 5000 54 211 8500 175 2006 9 000 300 3 465 500 988 800 157 300
2007 3000 62 580 10 300 3 2007 9 601 100 2 695 700 633 500 124 800
2008 2000 75 767 7300 4 2008 12 642 900 3 335 900 861 100 112 200
2009 2181 64 740 2600 216 2009 12 889 200 3 506 800 930 100 101 900
2010 0 153 473 3600 1878 2010 12 773 300 4 345 300 423 000 67 600
2011 185 155 513 2600 1850 2011 14 164 500 4 246 300 368 300 19 800
2012 10 142 329 2500 1280 2012 15 409 500 4 870 160 489 000 73 600
200312 1838 96 160 4305 564 200312 11 040 890 3 493 846 695 570 93 610

Italicized 5- or 10-year means. Italicized 5- or 10-year means.

Source: Food and Agriculture Organization of the United Nations (FAO). Source: Food and Agriculture Organization of the United Nations (FAO).

Table 4 Area harvested (ha) for major oilseeds grown in North Production
America
Canada has produced on an average about 11 Mt canola annu-
Year Canola Soybean Linseed Sunflower ally during the period of 200312 (Table 5). Canola produc-
tion in Canada has varied by a factor of about 2 during the
196367 479 195 13 922 433 1 817 502 60 922 2000s (Table 5). This variation resulted from the increase in
196872 1 276 286 17 560 138 1 676 065 209 421 harvested area as shown in Table 1 and about a 25% increase
197377 1 316 810 22 146 124 1 114 375 511 051
in yield with the introduction of hybrid cultivars. The average
197882 2 300 024 27 904 277 912 904 1 779 605
yield of canola in Canada has reached to 1.78 tonnes ha1 in
198387 2 699 707 25 566 435 878 543 1 134 925
198892 3 100 057 24 303 683 601 899 894 054 the recent years as compared to 1.35 tonnes ha1 in 1990s.
199397 4 857 249 26 283 954 740 011 1 243 165 The United States has produced on an average
19982002 5 172 135 30 302 314 893 539 1 233 620 754 943 tonnes of canola annually during the period of
2003 5 122 895 30 446 790 967 708 994 828 200312 (Table 6). Canola production in this country has
2004 5 206 240 31 192 400 727 697 747 915 increased by about 1.5-fold in recent years when compared
2005 5 631 034 30 096 036 1 120 480 1 127 062 with production in the 1990s (Table 6). The average yield of
2006 5 656 793 31 446 091 1 104 097 793 375 canola in the United States has reached 1.66 tonnes ha1, sim-
2007 6 798 670 27 193 325 675 537 893 139 ilar to the Canadian average yield.
2008 6 896 560 31 493 821 770 095 1 038 444
Mexico has produced on an average 2746 tonnes canola
2009 6 845 261 32 355 020 752 970 854 276
annually during the period of 200312 (Table 7). The produc-
2010 7 428 180 32 633 573 526 060 811 588
2011 7 894 105 31 554 323 345 810 605 610 tion of this crop in Mexico has reduced to almost half in the
2012 9 080 470 32 609 259 498 480 786 310 2000s when compared with 1970s (Table 7). The average yield
200312 6 656 021 31 102 064 748 893 865 255 of canola in Mexico has been 1.49 tonnes ha1, slightly lower
than Canada and the United States.
Italicized 5- or 10-year means. For the entire North American continent, average annual
Source: Food and Agriculture Organization of the United Nations (FAO). canola production was 11.8 Mt during the period of 200312
(Table 8); this is about 22% of the worlds production of
Brassica oilseeds. In North America, there has been a sharp
arable land) annually during the period of 200312 (Table 4); increase in production in the last 10-year period mostly
this production area constitutes about 22% of the worlds reflecting Canadian production. Canada has produced
Brassica oilseeds crop acreage. The canola production area in 93.6%, the United States 6.4%, and Mexico <0.3% of the
this continent has doubled in the recent years when compared North American canola during the period of 200312. In
with the production area in the 1990s (Table 4). Of the total terms of global production, North America has produced
North American canola harvested area during the period of about 22% of the worlds Brassica oilseeds during the period
200312, Canadas share was 93.1%, the United States 6.8%, of 200312. The share of world canola for Canada, the United
and Mexico <0.5%. States, and Mexico in the last 5 year average is 22.0%, 1.4%,
404 GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America

Table 6 Production (tonnes) for major oilseeds grown in the United Table 8 Production (tonnes) for major oilseeds grown in North
States America

Year Canola Soybean Linseed Sunflower Year Canola Soybean Linseed Sunflower

196367 0 22 592 554 682 119 40 319 196367 433 357 22 874 598 1 215 535 55 629
196872 0 31 646 150 626 942 155 861 196872 1 263 433 32 206 247 1 339 214 205 349
197377 0 40 105 814 344 966 596 032 197377 1 409 662 40 979 377 804 955 635 980
197882 0 55 071 062 233 298 2 256 880 197882 2 700 858 56 278 439 848 497 2 407 590
198387 2450 51 579 882 209 340 1 395 300 198387 3 433 783 53 326 233 967 827 1 461 224
198892 57 918 52 119 203 81 928 1 089 000 198892 3 816 190 54 051 454 628 429 1 176 857
199397 242 190 63 292 800 64 184 1 692 246 199397 6 364 140 65 843 445 952 010 1 769 907
19982002 769 017 75 110 850 247 081 1 726 263 19982002 7 410 317 77 667 860 1 068 078 1 849 665
2003 686 462 66 781 392 267 117 1 208 930 2003 7 458 662 69 180 698 1 022 477 1 351 385
2004 612 533 85 013 945 263 358 929 690 2004 8 288 133 88 191 192 782 258 982 122
2005 718 484 83 504 928 500 273 1 822 151 2005 10 206 784 86 847 763 1 491 833 1 906 582
2006 632 948 86 998 898 279 894 972 330 2006 9 641 248 90 545 511 1 274 694 1 129 808
2007 649 519 72 857 739 149 764 1 301 298 2007 10 254 619 75 641 810 790 464 1 426 105
2008 655 610 80 748 700 145 192 1 552 570 2008 13 302 510 84 237 622 1 011 392 1 664 775
2009 669 340 91 417 300 188 550 1 377 320 2009 13 561 821 95 045 000 1 120 450 1 479 552
2010 1 113 390 90 605 460 230 030 1 240 830 2010 13 886 690 95 118 425 655 530 1 312 227
2011 698 910 84 191 930 70 890 924 550 2011 14 863 440 88 643 464 440 990 947 176
2012 1 112 230 82 054 800 146 360 1 263 570 2012 16 521 880 87 172 460 637 060 1 338 478
200312 754 943 82 417 509 224 143 1 259 324 200312 11 798 579 86 062 395 922 715 1 353 821

Italicized 5- or 10-year means. Italicized 5- or 10-year means.

Source: Food and Agriculture Organization of the United Nations (FAO). Source: Food and Agriculture Organization of the United Nations (FAO).

Table 7 Production (tonnes) for major oilseeds grown in Mexico Table 9 Canada 10-year averages for disposition and consumption
(tonnes) 200009
Year Canola Soybean Linseed Sunflower
Crop Canola Soybean Linseed Sunflower
196367 3800 80 042 15 261 0
196872 5600 281 833 22 970 8376 Production 8 609 150 2 815 060 695 570 115 560
197377 5800 498 804 16 488 3056 Imports 200 832 479 988 13 000 21 823
197882 3600 543 900 5680 7730 Stock change 84 000 4000 1850
198387 1013 767 651 547 8484 Exports 4 799 534 1 242 257 530 570 82 127
198892 152 622 531 141 2757 Domestic supply 3 926 448 2 056 791 178 000 57 106
199397 1150 290 105 46 1321 Feed 288 376 336 197 5142
19982002 4400 118 730 258 562 Seed 45 593 122 440 30 260 954
2003 1000 126 006 960 155 Waste 266 662 27 000
2004 2000 133 347 2000 232 Processing 3 280 738 1 549 250 51 010
2005 5000 187 235 960 31 Food 45 080 21 804 8900
2006 8000 81 113 6000 178 Other uses 131 720
2007 4000 88 371 7200 7
2008 4000 153 022 5100 5 Statistics Canada, 2014.
2009 3281 120 900 1800 332 Source: Food and Agriculture Organization of the United Nations (FAO).
2010 0 167 665 2500 3797
2011 30 205 234 1800 2826
2012 150 247 500 1700 1308 canola used domestically. About 85% of the canola used domes-
200312 2746 151 039 3002 887 tically in Canada was used for processing and food manufacture,
7% for feed, and small amount for planting seed (Table 9).
Italicized 5- or 10-year means.
Source: Food and Agriculture Organization of the United Nations (FAO). Of the average annual 737 320 tonnes canola produced in
the United States and the 536 768 tonnes imported each year
during 2000s, 965 542 tonnes was used domestically and
and 0.003%, respectively. Currently, Canada ranks in the top
309 171 tonnes was exported (Table 10). Of the total domestic
five countries of the world for canola production.
supply in the United States, almost the whole amount was used
for processing and food manufacture (Table 10) and a small
amount for planting seed.
Disposition and Consumption
Mexicos domestic supply of canola is heavily dependent on
During 200009, of the average annual 8.6 Mt canola produced imports. During 200009, the country imported on an average
in Canada and the 200 832 tonnes imported, 3.9 Mt was used 1.1 Mt annually while producing only 5017 tonnes per year
domestically and 4.8 Mt was exported (Table 9). Thus, (Table 11). About 99% of the Mexicos domestic supply was
Canadian production was more than double the quantity of used for processing and food manufacture (Table 11).
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America 405

Table 10 US 10-year averages for disposition and consumption Soybean

(tonnes) 200009
Introduction
Crop Canola Soybean Sunflower
Soybean (Glycine max [L.] Merr.) was domesticated as a forage
Production 737 320 79 605 515 1 343 461 crop in China about 3000 BC from its wild progenitor, Glycine
Imports 536 768 215 039 71 636 soja. It became a commercially viable oilseed crop in the 1940s
Stock change 625 1 420 000 21 070 due to the development of shatter and disease-resistant new
Exports 309 171 29 835 045 147 185 cultivars and the need for a domestic source of edible oil and
Domestic supply 965 542 51 405 509 1 246 842 protein for feed while trade was disrupted during World War II.
Feed 1 272 200 353 400 There are only spring growth habit forms. By origin, the soy-
Seed 4159 2 433 077 11 042
bean progenitor is a subtropical plant, the cultivated soybean
Waste 17 177 1 736 721
has wide adaptation to various climatic and soil conditions. As
Processing 897 800 45 952 100 625 100
Food 72 469 11 413 265 400 a photoperiod-sensitive short-day plant, it is divided by lati-
Other uses 49 830 tude into 13 maturity groups (00010). While soybean has
been grown in China as a food crop for thousands of years, it
Source: Food and Agriculture Organization of the United Nations (FAO). is grown throughout the world and used primarily as a source
of edible vegetable oil and high-protein meal for animal feed
(Table 9). Recently, soybean has also been used as a source of
Table 11 Mexico 10-year averages for disposition and consumption biodiesel and bioproducts for the manufacturing industry such
(tonnes) 200009
as the automotive sector. Small quantities of soybeans are also
Crop Canola Soybean Sunflower used directly as animal feed, food (tofu, miso, natto,
soymilk, and soy sauce), and planting seed.
Production 5017 120 053 175
Imports 1 087 017 3 859 942 25 472
Stock change 680 0 700
Area Harvested
Exports 440 871 42 Canada has harvested soybean from about 1.3 Mha (10 year
Domestic supply 1 090 914 3 979 124 26 304 average, 200312, 2.9% of Canadian arable land) annually
Feed 867 170
(Table 1). The area of soybean harvested in Canada has varied
Seed 23 3365
by a factor of 3 during the 1990s, from about 555 000 ha to
Waste 11 026 119 359
Processing 1 072 080 2 185 877 23 790 over 1.67 Mha (Table 1). Until the mid-1990s, the majority of
Food 7785 1353 2514 soybean production area in Canada was in southwestern
Other uses 802 000 Ontario, whereas today, about 70% of soybean is grown in
Ontario with the remainder being grown in Manitoba, Quebec,
Source: Food and Agriculture Organization of the United Nations (FAO). and a small amount in Prince Edward Island.
The United States has harvested soybean from about
29.7 Mha (16.2% of American arable land) annually during
Table 12 North America 10-year averages for disposition and the period of 200312 (Table 2). The area of soybean harvested
consumption (tonnes) 200009 in the United States has increased since the 1990s, beginning
the decade at 23.6 Mha and ending it in 2012 at about
Crop Canola Soybean Sunflower
30.8 Mha (Table 2). The vast majority of soybean production
Production 9 351 487 82 540 628 1 459 196 area in the United States is in the midwestern US corn belt
Imports 1 824 617 4 554 969 118 931 states of Iowa, Illinois, Nebraska, Minnesota, Indiana, Kansas,
Stock change 84 055 1 424 000 18 520 Wisconsin, and South Dakota and the lower Mississippi Delta.
Exports 5 109 144 31 078 173 229 354 Mexico is a very small soybean producer, harvesting soy-
Domestic supply 5 982 905 57 441 424 1 330 252 bean from only 96 160 ha (0.4% of Mexican arable land)
Feed 288 376 2 475 567 358 542 annually during the period of 200312 (Table 3). The area of
Seed 49 775 2 558 882 11 995
soybean harvested in Mexico has varied widely during the
Waste 294 865 1 883 080
1960s to 2000s, from a low of 41 000 ha to a high of
Processing 5 250 618 49 687 227 699 900
Food 125 334 34 570 267 914 315 000 ha, with a trend toward decreased harvested area dur-
Other uses 49 830 802 000 ing the 1990s, which was replaced by an increasing trend in the
late 2000s (Table 3). This variation in harvested area is typical
Source: Food and Agriculture Organization of the United Nations (FAO). for any small-volume crop. Soybean production occurs in
central Mexico.
For the entire North American continent, soybean was har-
Of the average 9.4 Mt canola produced annually in North vested from an average 31.1 Mha (12.1% of North American
America during the period of 200009, 6.0 Mt was used arable land) annually during the period of 200312 (Table 4).
domestically (Table 12). About 90% of the canola used domes- The area harvested has varied slightly since 1990s, from a low
tically in North America was used for processing and food of 24.0 Mha to a high of 32.6 Mha (Table 4). Canadas average
manufacture (Table 12). A small amount was used directly soybean harvested area was 4.6%, the United States 94.9%,
for feed and planting seed as well. and Mexico <0.5% of the North American harvested area.
406 GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America

Production was used domestically and 871 tonnes was exported during the
period of 200009 (Table 11). Approximately 55% of the
Canada has produced on an average 3.5 Mt of soybean annu-
soybean used domestically in Mexico was used for food pro-
ally during the period of 200312 (Table 5). The production of
cessing (Table 11). Another 20% of the soybean domestic
soybean in Canada has varied by a factor of almost 3 during the
supply was used as animal feed and 20% was used in other
1990s and 2000s (Table 5), paralleling the variation in har-
uses. Minor amounts were used for planting seed and food
vested area for soybean in Canada shown in Table 1. The
(<3%).
average yield of soybean in Canada has been 2.8 tonnes ha1.
Of the average annual 82.5 Mt of soybean produced in
The United States has produced on an average 82.4 Mt of
North America and the 4.5 Mt imported, 57.4 Mt was used
soybean annually during the period of 200312 (Table 6).
domestically, and 31.1 Mt was exported during the period of
Soybean production has increased by more than 40% in the
200009 (Table 12). Over 60% of the soybean used domesti-
United States during the 1990s, from 52.1 Mt at the beginning
cally in North America was used for food processing (Table 12).
of the 1990s to 90.6 Mt at the end of the 2000s (Table 6). The
Small amounts were used for feed, seed, food, and other uses
average yield of soybean in the United States has been
(<8%).
2.9 tonnes ha1, similar to the Canadian average yield.
Mexico has produced on an average 151 000 tonnes of soy-
bean annually during the period of 200312 (Table 7). The
Linseed (Flax)
production of soybean in Mexico has varied by a factor of
8 during the 1990s, from a low of 81 000 tonnes early 1960s Introduction
to a high of 622 000 tonnes in early 1990s (Table 7). This
Linseed (Linum usitatissimum), also known as flax in North
variation in production is typical for any medium volume
America, is one of the earliest cultivated field crops, initially
crop. The average yield of soybean in Mexico during the
grown for its fiber. In North America, in the last two centuries,
1990s has been 1.7 tonnes ha1, lower than the soybean yields
linseed has been grown primarily for its oil. It is a spring
seen for Canada and the United States. Soybeans are grown in
annual, adapted to a wide range of soil and climatic conditions
Mexico under water-limiting rain-fed conditions.
in the cool temperate zones of the northern hemisphere. Lin-
For the entire North American continent, average annual
seed in North America is used primarily to produce industrial
soybean production during the period of 200312 was
oil and animal feed meal. Linseed oil is a drying oil used in
86.1 Mt (Table 8). There has been a year-to-year variation in
paints and varnishes and for the manufacture of linoleum
production, with a trend toward increased production, not
flooring. A trend toward bioproducts has increased this market
only mostly in the United States in recent years but also in
in the 2000s. Because linseed oil contains alpha-linolenic acid
Canada. In terms of North American production proportions,
(ALA), an essential omega-3 fatty acid, it has a variety of uses.
Canada has produced an average 4.6%, the United States
Whole linseed is fed to poultry to produce omega-3 fatty acid-
94.9%, and Mexico <0.5% of the soybean produced in
enriched eggs. Linseed meal is a component of rations fed to
North America. In terms of global soybean production,
pets, poultry, and animal. Small quantities of linseed are also
North America has produced on an average 33.0% of the
used in food products (breads and breakfast cereals), but a
soybean produced in the world. The share of world soybean
significant number of products are being developed using
production for Canada, the United States, and Mexico is
linseed, in response to the variety of health benefits, including
1.0%, 32.5%, and <0.2%, respectively. The United States
reducing heart disease and cancer risk. Additional products
is the top soybean producer of the world; however, it falls
have been created based on the health and nutrition claims
behind Brazil and Argentina, which represent almost a half
of linseed oil, lignans, and antioxidant flavonoids. Small quan-
(49%) of world production combined.
tities of linseed are used for (re)planting seed.

Disposition and Consumption Area Harvested

Of the average annual 2.8 Mt of soybean produced and
480 000 tonnes imported by Canada, 2.1 Mt was used domes-
tically and 1 242 000 tonnes was exported during the period of
200009 (Table 9). Approximately 55% of the soybean used
domestically in Canada was used for food processing, mainly
edible oil production (Table 9). Another 12% of the soybean
domestic supply was used as animal feed. Minor amounts were
used for planting seed and food (5%).
Of the average annual 79.6 Mt of soybean produced in the
United States and the 215000 tonnes of soybean imported,
51.4 Mt was used domestically and 29.8 Mt was exported during
the period of 200009 (Table 10). Approximately 64% of the
soybean used domestically in the United States was used for food
processing, mainly edible oil production (Table 10). Small
amounts were used for feed, planting seed, and food (<5%).
Of the average annual 120 000 tonnes of soybean produced
in Mexico and the 3.9 Mt of soybean imported, about 3.9 Mt
Canada has harvested linseed from an average of 552 000 ha
(1.25% of Canadian arable land) annually between 2003 and
2012 (Table 1). The area of linseed harvested in Canada has
varied widely, from a low of about 273 000 ha to a high of
785 000 ha (Table 1). The vast majority of linseed production
area in Canada is in the Prairie Provinces (Alberta, Saskatche-
wan, and Manitoba).
The United States has harvested linseed from about
192 000 ha (0.14% of American arable land) annually, aver-
aged between 2003 and 2012 (Table 2). The area of linseed
harvested in the United States has varied widely during the
2000s, from a low of about 70 000 ha to a high of 386 000 ha
(Table 2). The vast majority of linseed production area in the
United States is in the north-central states of North and South
Dakota, Minnesota, and Montana.
Only a small amount of linseed is grown in Mexico, aver-
aging 4300 ha between 2003 and 2013.
 GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America
For the entire North American continent, linseed was har-
vested from an average 748 000 ha (0.32% of North American
arable land) annually between 2003 and 2012 (Table 4). The
area harvested has varied widely during the 2000s (Table 4).
Canadas average linseed harvested area was 73.7%, United
States 25.6%, and Mexico 0.6% of the total North American
linseed harvested area.
Production
Canada has produced on an average 695 570 tonnes of lin-
seed annually between 2003 and 2012 (Table 5). The pro-
duction of linseed in Canada has varied widely during the
2000s, from a low of 368 000 tonnes to a high of 0.99 million
tonnes (Mt) (Table 5), paralleling the widely varying produc-
tion area for linseed harvested in Canada as shown in Table 1.
The average yield of linseed in Canada between 2003 and
2012 was 1.26 tonnes ha1.
The United States has produced on an average
224 000 tonnes of linseed annually (Table 6). The production
of linseed in the United States has varied widely during the
2000s, from a low of 71 000 tonnes to a high of 500 000 tonnes
(Table 6), paralleling the widely varying production area for
linseed harvested in the United States shown in Table 2. The
average yield of linseed in the United States has been
1.02 tonnes ha1, slightly lower than the Canadian average
yield.
For the entire North American continent, average annual
linseed production was 923 000 tonnes annually between 2003
and 2012 (Table 8). In terms of North American production
proportions, Canada has produced an average of 75%, United
States 24%, and Mexico 0.3% of the linseed produced in North
America. In terms of global linseed production, North America
has produced on an average 43.6% of all the linseed produced in
the world. The share of world linseed production for Canada and
the United States is 32.8% and 10.5%, respectively. Canada was
the top linseed producer in the world in the 2000s, dropping
from 34.8% in the 1990s due to the identification of low-level
presence (LLP) of genetically engineered flax seeds.
Disposition and Consumption
Canada is a net exporter of linseed. Of the average annual
695 570 tonnes of linseed produced in Canada and the
13 000 tonnes imported into Canada, 178 000 t was used
domestically and 530 570 tonnes was exported, primarily for
industrial oil uses (Table 9). Of the linseed used domestically
in Canada, only small amounts of linseed (<5%) were used
for food, while 74% was used for other uses combined (seed
crushed for industrial oil and meal production for feed), and
17% was used for planting seed (Table 9). An average of
304 000 tonnes was retained on farm, for either seed or tem-
porary storage (Statistics Canada 2014). Canada is a major
exporter of linseed to the world (530 570 tonnes), with sig-
nificant exports to Europe, the United States, Japan, South
Korea, and China. Canada currently supplies nearly 60% of
the linseed used in the United States.
The United States is a net importer of linseed but exports
80% linseed to either Belgium or Canada. Because linseed is a
relatively minor bioproduct crop, internal disposition of flax is
not tracked by USDA or FAO.
407
Sunflower
Introduction
Sunflower (Helianthus annuus) was domesticated as a food crop
in North America, perhaps as early as 3000 BC. This was
introduced to Europe in the 1600s and was successfully devel-
oped as an oil crop in Russia in the early 1800s. Russian plant
breeders were able to increase the oil content in sunflower seed
from less than 30% to over 50% the major factor permitting
the development of sunflower as oil crop for the temperate
areas of the world.
Sunflower is used primarily for edible vegetable oil and the
seed meal is used for animal feed (Table 10). Moderate quan-
tities of sunflower are directly used for animal feed and food
manufacture (confectionary and bakery products) (Table 10).
Small amounts are used for planting seed (<5%).
Area Harvested
Canada has been growing sunflower commercially since the
early 1940s; however, the production area remains small. Dur-
ing the period of 200312, Canada on an average has har-
vested sunflower from about 62 500 ha (about 0.1% of
Canadian arable land) annually (Table 1). The area of sun-
flower harvested in Canada varied greatly during the years
since the 1960s from a low of <15 000 ha to a high of
>100 000 ha (Table 1), which is typical for a small-volume
crop. The vast majority of sunflower production area in
Canada is in the Prairie Province of Manitoba.
The United States on an average has harvested sunflower
from about 0.8 Mha (0.5% of American arable land) annually
during the period of 200312 (Table 2). This is about a 15-fold
increase in the production area when compared with the pro-
duction area in 1960s; this is, however, about a 30% reduction
when compared with the 1990s. The vast majority of sunflower
production area in the United States is in the central Plains
states (from Dakotas to Texas, with North Dakota being the
predominant production area state).
Mexico has been growing sunflower since early 1970s. Dur-
ing the period of 200312 the country on an average has
harvested sunflower from only 564 ha (0.002% of Mexican
arable land) annually (Table 3); this is <10% of the produc-
tion area when compared with the acreage of this crop in the
1970s and 1980s. The area of sunflower harvested in Mexico
has varied widely since 1970s; the crop almost vanished in
mid-2000s (Table 3). The greater part of the sunflower pro-
duction area is in central Mexico.
For the entire North American continent, sunflower was
harvested from an average 0.9 Mha (0.4% of North American
arable land) annually during the period of 200312 (Table 4);
this constitutes about 3.6% of the total sunflower production
area in the world. The area harvested in North America has
varied greatly in the last 50 years (Table 4). Of the total
sunflower harvested area in this continent during 200312,
Canadas share is 7.2%, the United States 92.7%, and Mexico
only 0.07%.
Production
Canada has produced on an average 93 610 tonnes of sun-
flower annually during the period of 200312 (Table 5). The
408 GRAINS AROUND THE WORLD | Grain Production and Consumption: Oilseeds in North America
production of sunflower in Canada has varied greatly from a
low of <20 000 tonnes to a high of >150 000 tonnes (Table 1)
reflecting the variation in harvested area. The average yield of
sunflower in Canada has been 1.5 tonnes ha1.
The United States has produced on an average 1.3 Mt of
sunflower annually during the period of 200312 (Table 6)
with an average yield of 1.6 tonnes ha1 (similar to the
Canadian average yield). Compared to the 1990s, the country
has produced about 20% less sunflower in the last 10 years.
Mexico has produced on an average 887 tonnes of sun-
flower annually during the period of 200312 (Table 7). The
production of sunflower in this country has varied greatly; this
is primarily due to the variation in production area (Table 1).
The average yield of sunflower in Mexico has been
1.6 tonnes ha1, very similar to the yields for Canada and the
United States.
For the entire North American continent, average annual
sunflower production was 1.4 Mt during the period of
200312 (Table 8). This is about 23% reduction in production
when compared with the production in the 1990s. In terms of
North American production proportions, Canada has pro-
duced 6.9%, the United States 93.0%, and Mexico <0.1% of
the total sunflower produced in North America during
200312. In terms of world sunflower production during
200312, North America has produced 4.2% of all the sun-
flowers produced in the world (Canadas and Mexicos share is
<0.4% and the United Statess share is 3.9%).
Disposition and Consumption
Of the average annual 115 560 tonnes of sunflower produced
and the 21 823 tonnes of sunflower imported in Canada dur-
ing the period of 200009, 57 106 tonnes was used domesti-
cally and 82 127 tonnes was exported (Table 9). About 89% of
the sunflowers used domestically in Canada were utilized in
processing and food manufacture, 9% was used as animal feed,
and a small amount was used as planting seed (2%) (Table 9).
Of the average annual 1.3 Mt of sunflower produced and
the 71 636 tonnes of sunflower imported in the United States
in the 2000s, 1.2 Mt was used domestically and 147 185 tonnes
was exported (Table 10). About 70% of the sunflowers used
domestically in the United States were utilized in processing
and food manufacture, 28% were used as animal feed, and 1%
were used for planting seed (Table 10).
Mexicos average annual domestic supply of sunflower in
the 2000s was about 26 300 tonnes of which only 1% was
produced within the country and the rest being imported
(25 472 t) (Table 11). All domestic supply of the sunflower in
Mexico was used in processing and food manufacture
(Table 11).
Of the average annual 1.5 Mt of sunflower produced and
the 118 931 tonnes of sunflower imported in the North Amer-
ican countries during the period of 200009, about 84% of this
supply was used domestically (Table 12). Over 70% of the
sunflower used domestically in North America was used for
processing and food manufacture, 27% was used as animal
feed, and 1% was used for planting seed (Table 12).
Exercises for Revision
Describe the contrasts in the origins and cultivation of these
major oilseed species in North America.
Explain the role of plant breeding in the improvement of
these crop species.
What features distinguish oilseed production, quality, and
consumption in your region from these factors in North
America (or choose one of the North American countries)?
Compare and contrast the utilization of the products of
these four major oilseeds.
Exercises for Readers to Explore the Topic Further
Flax is a minor but significant crop in North America, where
it is primarily grown for its seed. Find out about the possi-
ble use of the traditional product of flax as a fiber.
Find out the differences in composition and properties
between the oils obtained from these four oilseed species.
Find out the differences in composition and properties
between the residues (meal) left after oil removal obtained
from these four oilseed species.
Obtain details of oilseed-production areas of North Amer-
ica for each of the species. How similar (or different) are the
respective growing sites in each of these countries?
See also: Agronomy of Grain Growing: Canola: Agronomy;
Soybean: Agronomy; Breeding of Grains: Canola/Rapeseed:
Genetics and Breeding; Soybean: Germplasm, Breeding, and Genetics;
Fats: Healthy Fats and Oils; Lipid Chemistry; Grain Composition
and Analysis: The Composition of Food Grains and Grain-Based
Products; Grain Harvest, Storage and Transport: Canola:
Harvest, Transport, and Storage; Grains Around the World: Grain
Production and Consumption: Cereal Grains in North America; Non-
food Products from Grains: Cereal Grains as Animal Feed; Oilseed
and Legume Processing: Canola: Processing; Soybean:
Processing; Soybean: Soy Concentrates and Isolates; The Oilseeds:
Canola: Overview; Linseed: Overview; Oilseeds: Overview; Soybean:
Overview; Sunflower: Overview.
Further Reading
Robbelen G, Downey RK, and Ashri A (eds.) (1989) Oil Crops of the World, p. 554.
Toronto: McGraw-Hill.
Slinkard AE and Knott DR (eds.) (1995) Harvest of Gold: The History of Field Crop
Breeding in Canada, p. 367. Saskatoon, Saskatchewan: University Extension Press/
University of Saskatchewan.
Relevant Websites
http://www.canolacouncil.org/ Canola Council of Canada.
http://www.canadasunflower.com/ National Sunflower Association of Canada (NSAC).
http://faostat3.fao.org/faostat-gateway/go/to/download/Q/*/E Food and Agriculture
Organization of the United Nations.
http://www.sunflowernsa.com/ National Sunflower Association.
http://www.statcan.gc.ca/start-debut-eng.html Statistics Canada.
http://www.uscanola.com/ U.S. Canola Association.
http://usda.gov United States Department of Agriculture (USDA).
 Grain Production and Consumption in Oceania: Australia and Pacific Countries
CW Wrigley, University of Queensland, Brisbane, QLD, Australia
RJ French, University of Western Australia, Crawley, WA, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Food and Agriculture Organization lists 26 countries in the
Oceania region the South Pacific including the main
grain producers Australia and New Zealand.
Maize and rice are produced in tropical countries such as
New Caledonia, the Solomon Islands, New Caledonia,
Vanuatu, and Fiji.
The introduction of cereal grains to Australia since 1788 has
been difficult with inappropriate varieties and upside-
down seasons.
Australia produced the worlds first wheat breedercereal
chemist combination (FarrerGuthrie) in the 1890s.
Farrer and other breeders have greatly improved the yields
and grain quality of Australian wheats.
Grain yields differ considerably between grain species and
between Australia and New Zealand.
Grain yields are expected to increase as atmospheric levels
of CO2 rise in coming decades; the protein content will
decrease as a result.
Learning Objective
To extend knowledge about the production and utilization
of grains around the world by learning about the countries
in the Oceania region
Introduction
The Oceania region includes the many islands of the southern
Pacific Ocean, including Australia (classed as both an island
and a continent), New Zealand (two major islands), Papua
New Guinea, and many smaller islands. Those smaller islands
that produce significant amounts of grain are listed in Table 1.
Others include the Cook Islands, French Polynesia, Nauru,
Samoa, and Tonga, most of them in the tropics. The Food
and Agriculture Organization (FAO) lists 26 countries in Oce-
ania in its statistics of agricultural products. Of these countries,
the main grain producers are Australia, New Zealand, and
Papua New Guinea (Table 2). Australia produces by far the
greatest volume and diversity of grains, being also by far the
largest in area, although much of its land area is not suited to
grain production (Figure 1).
None of the grains of economic significance are native to
any of the countries of Oceania. These grains have been intro-
duced by the first European colonists whose arrival in the
southern Pacific is only a little over two centuries ago. The
first settlers in Eastern Australia (the Sydney of 1788) found
themselves trying to grow wheat under conditions in the
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00049-8
southern hemisphere that were completely different from
those of their homeland, England.
The first wheat crop, grown at the current site of Sydneys
Royal Botanic Gardens, yielded little more than the few
bushels that were sown. In contrast to those early days,
Australia is now a major exporter of wheat and of many other
grains. Wheat and wool have been important for Australias
(early) prosperity. Their importance has been acknowledged
on the first bank notes issued in 1966 at the commencement of
decimal currency (Figure 2).
New Zealand was colonized at a time and under circum-
stances similar to those of Australia. Grain production in New
Zealand is now also significant (Table 2), but generally not to
the extent of being involved significantly in grain export. In
contrast, the many nations of the Pacific Islands are importers
of grain and flour products. For these reasons, much of this
article relates to grain production in Australia.
Historical Perspective
In its early decades, the Australian grain-growing industry fluc-
tuated from underproduction through self-sufficiency to occa-
sional periods of excess (Table 3). Wheat was the main grain
grown, being needed for bread making. Farmers had to contest
dry conditions, poor soil in some of the areas cultivated ini-
tially, attack by the rust pathogen, the reversal of seasons from
their previous experience in the northern hemisphere, and
varieties that were poorly adapted to the new continent.
The problem of inappropriate wheat varieties was realized
in the latter half of the nineteenth century by far-sighted
farmers, by staff of experimental farms, and by an amateur,
William Farrer, whose profession as a surveyor took him into
the heart of the wheat-growing regions. Whereas farmers
attempted to develop improved varieties by selecting seed
from plants that looked better than the rest of the crop, Farrer
set about the (then) new technique of crossbreeding to increase
the genetic diversity from which to select better genotypes.
Figure 3 is an artists impression of Farrer among his small
plots of crossbred wheats. The tall plants contrast with the
much shorter wheats of more recent times.
An example of the selection method is given in Table 3: the
extension of wheat growing into drier regions was attributed to
the selection of Purple Straw (an earlier-maturing wheat)
from a field of Red Straw by a South Australian farmer in
about 1860, just before the introduction of crossbreeding. The
recent use of crossbreeding to produce new wheat varieties is
shown in Figure 4. In most cases, crosses have been made, but
there are still a few new varieties that have arisen as selections
from an already existing variety, such as the selection of
Meering from the former wheat Condor, which was presum-
ably released as a mixture of a few genotypes.
409
410 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries

Table 1 Production, in kilotonnes, of grains according to categories improvement of grain quality for all grain species. A world-first
in crop year 2012 for countries of Oceania where there is significant innovation of that time (1890s) was the development of a very
production small-scale flour mill, suited to evaluating the milling quality
of the small samples of grain coming from Farrers wheat
Country Cereal grains Oilseeds Pulses
breeding (Figure 5). Subsequent years brought the excess pro-
Australia 43 372 5291 2956 duction of wheat that justified Farrers vision of export to
Fiji 9 0.3 2.4 England, largely as a result of the development of new wheat
Micronesia 0.3 varieties (Table 3). These contributions by Farrer were recog-
New Caledonia 3 nized by the issue of $2 bank note when Australia changed to
New Zealand 1168 5 23 decimal currency in 1966 (Figure 2).
Papua New Guinea 16 1.4 4 However, the greatest expansion in wheat production
Solomon Islands 4 6
occurred in the second half of the twentieth century, with
Vanuatu 0.8 3
production expanding from about 2 million tonnes (Mt) in
Source: http://faostat.fao.org. 1940 to a high of about 25 Mt in 1999/2000. Nevertheless,
production has fluctuated on occasions due to drought. The
high of 2000 contrasts with the drought year of 2002/2003,
when the national wheat crop was only a little over 10 Mt, less
Table 2 Production, in kilotonnes, of grain crops for the 2012 crop
than half of the crops either side (about 24 Mt in both 2001/
year in Australia, New Zealand, and Papua New Guinea (PNG)
2002 and 2003/2004). Similar reductions in production were
Crop Australia New Zealand PNG experienced for all grains in the 2002/2003 drought year. More
recently, fluctuations have not been so great (Table 4) and total
Cereals wheat production has continued to grow, with the production
Barley 8221 439 of nearly 30 Mt in 2012.
Maize 451 211 10
Millet 45
Oats 1262 18
Phalaris, canary seed 6 Diversification of Markets for Australian Wheat
Rice, paddy 919 1.2
Rye 40 Further improvements in the quality of wheat varieties and in
Sorghum 2239 4.5 management practices through the early twentieth century
Triticale 285 extended opportunities for grain exports. Initially, these
Wheat 29 905 489 exports were to England, where the grain was used for tradi-
Oilseeds tional products, mainly bread. This situation changed dramat-
Cottonseed 1694 ically in the second half of the twentieth century. In recent
Linseed 7 2.4
decades, Australia has produced about four times more than its
Peanuts 25 1.4
domestic needs, so that 7080% of the wheat harvest is
Rapeseed 3427 2.7
Safflower 5 exported. Exported wheat goes to a wide range of countries,
Soybeans 86 including Egypt, Indonesia, Iran, Iraq, Japan, and China.
Sunflower seed 47 In the early days of this expansion, there was minimal
Pulses 4 diversification of the crop on the basis of grain quality. Fair
Beans (various) 475 average quality was the basis of assessment of quality for most
Chickpeas 673 of the crop, although the region of origin of the FAQ wheat
Lentils 463 3 provided a good indication of quality (Figure 6). The 1960s
Lupins 982 brought the realization that export markets for Australian
Peas 343 20
wheat included a diversity of foods that could then be classed
Vetches 20
as exotic (to Australians), such as Arabic flat breads in the
Gaps indicate insignificant production. Middle East and a range of noodles and steamed breads in
Source: http://faostat.fao.org. Asian countries. Furthermore, in countries such as Indonesia,
even the manufacture of leavened bread involved non-
traditional methods that required wheat of quality different
Farrers use of crossbreeding was partly aimed at developing from the English market.
wheats with grain quality suited for export to England by Accordingly, experts were brought to cereal chemistry lab-
combining the dough quality of Canadian Fife wheats with oratories in Australia from the new export markets to demon-
the drought tolerance of wheats introduced from countries strate the production of these exotic products. Figure 7 shows
such as India, Palestine, and South Africa. This vision led him an expert from the Middle East with experimental Arabic
to enlist the help of the agricultural chemist, Frederick Guthrie, pocket breads, from both normal-scale baking and small-
to evaluate the milling and baking quality of the small samples scale baking. As a result of seeking this expertise, it was possible
of his many crossbred lines. for wheat breeders to select new varieties that would suit these
Thus, there was a start to the development of small-scale diverse quality requirements. The resulting research led to the
test methods and (probably) the worlds first breederchemist establishment of specifications for the wide range of wheat-
association, a combination that is now an integral part of the based products made around the world. With the publication
 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries 411

Australian wheat growing regions

Wheat terminals
Australian premium white wheat area
Predominantly Australian premium
white and hard wheat area
Predominantly hard wheat area

Northern
Territory

Queensland

Western Australia

South Australia
New South
Wales

Victoria

Tasmania

Figure 1 Wheat-growing areas of Australia. Reproduced from Australian Crop Report, 2007, Australian Bureau of Agricultural and Resource
Economics, Canberra.

Figure 2 The $2 bank note, introduced with Australias decimal currency in 1966, featured William Farrer the wheat breeder responsible, in the period
18901910, for providing farmers with new varieties adapted to Australian conditions.

of these findings, advantages of the knowledge have flowed to
the importing countries involved and to other exporters.
Another form of diversification came with the challenge
presented to wheat growers by the sandy soils of Western
Australia, due to the lack of soil nitrogen. An efficient solution
to this problem was the introduction of grain-legume crops in
agronomic rotation with wheat. The ability of the legumes to
fix atmospheric nitrogen led to the provision of a natural
source of nitrogen fertilizer, plus the welcome harvest of
other grains, particularly lupins and various pea and bean
crops (Table 2). These pulses have only emerged as important
crops since the 1980s. Expansion of production has depended
significantly on the growth of export markets. Recent expan-
sions in wheat and pulse production have warranted the
enlargement of terminal storage facilities in Perth, Western
Australia (Figure 8).
Another form of agricultural diversification accompanied
the downturn in wheat prices during the 1980s. In addition to
growing grain legumes, a valuable crop was supplied in the
form of canola (rapeseed). Production of canola continued,
Table 3 Historical developments in the Australian wheat industry

I. Period of foundations 178892

1788 Eight acres is sown to wheat at Farm Cove, Sydney
1790 James Ruse grows bearded wheat on a few acres at Rose Hill, Parramatta, in Western Sydney
1792 About 200 acres is cultivated to wheat in the colony
II. A wheat market is established 17931824
1793 The colony is self-sufficient in wheat and maize
1803 Rust destroys wheat crops in Sydneys Dundas Valley
1804 Abundant yields of Common Brown wheat in the Sydney district bring self-sufficiency again, but prices are low
1810 White Lammas and Red Lammas wheats from England and Scotland are cultivated, mainly in coastal regions. White Lammas was late-
maturing and poor in baking quality
1811 James Ruse makes himself a plow; previously most tillage was by hoe
1822 Various wheats are introduced by the newly established Royal Agricultural Society of New South Wales from England, South Africa, India, and
Egypt
III. Period of insufficient expansion 182555
1850 Red Straw and White Essex (a Lammas type) are introduced
IV. Period of declining yield 185596
1860 South Australias climate and geography favor quality-wheat production, and grain is exported to England
1860 Purple Straw (an early-maturing wheat) is selected, probably from a field of Red Straw by a South Australian farmer. This selection starts
an extension of wheat growing into drier regions
1880 Introduction of Ganz steel roller mills from Hungary alters quality requirements
1881 Du Toits wheat, early-maturing and rust-resisting, is introduced from South Africa by Dr. Schomburgk, director of the Adelaide Botanic
Gardens. From this introduction, James Ward, of Port Pirie, selects a rust-resisting high-yielding wheat named Wards Prolific
1882 Correspondence with The Australasian newspaper prompts the English-born surveyor W. J. Farrer to make plans for wheat improvement by
crossbreeding and selection, thereby going beyond the previous practice of selection only for wheat improvement
1889 Farrers first crosses are attempted at his property, Lambrigg, near Canberra. The variety Blounts Lambrigg is selected from crossbred
material provided by A. E. Blount of Colorado, the United States
1890 Disastrous wheat losses, due to rust, prompt the first rust-in-wheat conference to be convened. Farrers letter to the conference calls for
crossbreeding as a means of improving both rust resistance and grain quality
1893 F. B. Guthrie, recently appointed chemist with the Department of Agriculture of the Colony of NSW, devises small-scale tests of milling and
baking quality for selecting suitable parents and crossbreds from Farrers breeding program. Improvement of quality is thereby made
possible
1893 Hugh Pye produces Improved Steinwedel at Dookie College, Victoria, by making the cross (Steinwedel  Purple Straw)  Steinwedel
1893 A date suggested as the origin of cereal chemistry as a discipline, due to breederchemist collaborations in Australia and overseas
1895 Farrers Yandilla is the first of a new generation of wheats, designed to combine Fife quality with the drought resistance and earliness of
Indian wheats
V. Period of rapid expansion 18961930
1896 Local millers are forced to modify their machinery to suit imported North American Fife wheats due to local shortages. They are thus more
ready to accept Farrers stronger wheats, which they had previously rejected
1898 Farrer is appointed officially as wheat experimentalist to the NSW Department of Agriculture on an annual salary of 350
1900 Release of Bobs, reportedly obtained by crossing a selection from Blounts Lambrigg with Bald Skinless Barley by Farrer in 1896. It
became popular throughout Australia and represented a breakthrough in the production of strong wheat
1901 Export of 25 million bushels of wheat and flour
1901 Release of Federation, selected by Farrer from his cross in 1895 between Purple Straw and Yandilla. Though it fell short of his quality and
disease-resistance goals, its yield made it the most popular wheat in Australia from 1910 to 1925. It was also widely grown in the United
States, India, and other countries
1906 Death of Farrer and suspension for some years of quality-directed breeding
1911 NSW Strong White Wheat class is established to segregate Farrers strong wheats. Export of 64 million bushels of wheat and flour of all
classes. The contrast between this harvest and that of 10 years before is due largely to Farrer wheats such as Federation
1924 Release of Ghurka, bred by G. S. Gordon in 1916, becoming a leading Victorian variety in its own right and also a parent of many important
soft wheats
1929 The variety Bencubbin commences its phenomenal rise to popularity, not only in Western Australia, where it was bred by E. J. Limbourn, but
also throughout Australia
VI. International marketing expansion 193048
1936 The Gepp Royal Commission recommends that breeders should endeavor to produce new varieties that have the quality of strength
1945 Availability of Gabo wheat saves many farmers from ruin due to rust. It starts Australias reputation for high-protein baking-quality wheat
1946 Introduction of Insignia, selected for tiller survival, permits further extension of the wheat belt into drier areas
VII. Grain quality targeted to specific market needs 194970
1950 Segregation of specific truckloads of high-protein wheat of selected varieties as prime-wheat grades by the Prime Wheat Association, in
northern New South Wales
1956 Release of Dural, the first Australian-bred durum wheat
1957 Passage of the Wheat Research Act through federal parliament and the consequent formation of the Wheat (Industry) Research Council, later
to be replaced by the Grains R&D Corporation with responsibility for research on the wide range of agricultural grains
1960 Naming of Gamenya, derived from Gabo by I. A. Watson. Though bred in the east, its greatest impact was in Western Australia and as an
ideal noodle wheat
1960 Growing market awareness, involving an active acknowledgment that about three-quarters of the Australian wheat crop is exported for
exotic uses, leading to research initiatives on noodles, flat breads, and steamed bread
1967 Timgalen, issued by the University of Sydney, offers excellent quality with high-protein potential, disease resistance, and agronomic
qualities that earned it a continuing unrivaled position for prime-wheat production
1970 Introduction of near-infrared spectroscopy revolutionizes the analysis of protein and moisture in grains
 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries
Figure 3 Artists impression of the 1890s wheat breeder William Farrer
among his small plots of crossbred wheats.
tripling in the period 199699; cottonseed also increased with
the expansion of cotton production (Table 2).
Wheat
The Australian Wheat Board was given responsibility for mar-
keting the wheat crop since its formation in 1939. In July 1999,
its status changed from an Australian government statutory
marketing authority to a private company called AWB Limited.
Since then, there has been further diversification of marketing,
with the result of general deregulation.
Wheat yields in Australia are relatively low by world stan-
dards (Table 5) largely because production is mainly dryland,
with only a small extent of irrigation, reflecting Australias
character of being the worlds driest continent.
Wheat varieties registered in Australia are virtually all white-
grained, in contrast to the red-grained wheats that are common
in other wheat-producing countries. This policy direction,
taken in the very early days of the industry, has been recog-
nized as a marketing advantage. For many years, only white-
wheat varieties have been grown. In addition, the dry climate
of Australias wheat belt (mainly inland) ensures that the
grain produced is very dry. Furthermore, marketers and
413
bulk-handling corporations set rigid standards for the cleanli-
ness of export wheat.
Consequently, Australian wheat has regularly been mar-
keted as clean, white, and dry. These three characteristics
offer millers the triple promise of high flour extraction,
namely, the lack of nonmillable material (clean), the white
grain permitting a higher extraction rate before flour color is
compromised, and dry grain offering less moisture to be paid
for at the price of grain (if sold on as is weight).
Because of the high proportion of the Australian wheat crop
that is exported, marketing strategies must take into account
the wide range of products for which it is used. Several of these
are shown in Figure 9. The main products domestically are the
many conventional forms of leavened bread. This type of
baked product is in a minority for export uses, which are
more likely to include products such as various Arabic flat
breads, Chinese steamed breads, many types of noodles
(made from hexaploid wheats), and pasta (made from tetra-
ploid durum wheats). Durum wheats make up a small propor-
tion of the crop, but this proportion is increasing.
To meet the wide range of quality specifications for these
various products, there are many wheat grades with tight spec-
ifications. Varietal identity is an important part of the quality
specifications, with a small group of varieties of similar quality
type being permitted for each grade. Other specifications cover
aspects of the plumpness of the grain, absence of contaminants
and of insects, cleanliness, soundness, the absence of defects,
and, importantly, protein content. This approach to grading
and marketing is in great contrast to the FAQ era of the earlier
part of the twentieth century.
The maintenance of grain quality in the current range of
grades and classes requires strict systems for assessing grain
quality when it is delivered. It is common for grain to be
delivered directly from the field to the local silo (grain elevator
or storage site), although on-farm storage is becoming more
common. At the point of delivery, it is critical for the range of
quality attributes to be determined quickly, to allow for the
grain to be dumped into the storage cell appropriate to the
grade to which it is allocated. In addition, grain samples are
kept to represent the deliveries of each grower and to indicate
the composition of the storage cells. These samples can be
further analyzed subsequent to delivery. Identification of vari-
ety is also needed for the payment of royalties, based on
deliveries, according to plant breeders rights.
Rice
Rice production and export has been a significant industry for
Australia, but production has been reduced in recent seasons
by drought. Table 4 shows the low point of less than
200 000 tonnes of paddy rice in 2010, but production has
since risen again. It is grown within the irrigation areas of the
Murrumbidgee and Murray River systems in southern NSW,
with some further areas of production in Victoria, Western
Australia, Queensland, and the Northern Territory.
Farming operations are entirely mechanized. In the irrigation
area, sowing occurs in September and October into dry seedbeds.
Sometimes, sowing involves the aerial dispersal of pregerminated
seed into shallow water. After about 4 months, water is drained
414 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries

Raven
Aroona
WW15
2* Condor Meering
WW80 2* Brochis S Batavia
Banks
PWTH Janz
Cook
Winglen 4* Cunningham
3Ag3
Timgalen WW15 3* Sunco
AKMS 3Ag14
Sunvale
Madden 5*
SUN9E-27 4*
VPM1
Gabo Sunstate
2* Machete
PavonS Hartog 4*
Kite Amber mutant 2* Sunbrook
2* Blade
TPP Sonora 64
Spica
Yaqi54 BT-Schomburgk Suneca
MEC3 Ciano
Barunga
Sonora 64 Molineux Wilgoyne
Sabre Frame Gallo
Dagger
Wren
Insignia Spear Stiletto
RAC177 3* Gaboto
3* Dollarbird
Condor Vereanopolis Kalyansona

Heron Rosella Bluebird

Farro Lunga
WW33B WW33G Cocamba
4* Ouyen
TA3PNB3P TM56
AUS10894 Takari

Bencubbin Siete Cerros

Insignia Bodallin
Bokal
Gabo
6*
Mentana Gamenya
2*
Kenya 117a Eradu
Ciano
Wilgoyne Sunfield 51b
Gallo Cunderdin Perenjori
Zambesi
Noroeste Hyden 3*
Cranbrook Inia 66 Amery
Wren Shortim
Charter 2*
Lr21-SrX
Kenya C604 Jacup
Bencubbin Cadoux
Sword Mexico 2*
Centrifen Falcon-EMS
Madden
Gabo
3* Sharbarti
Khapstein Sonora 64 Kalannie
PI0522B
GluyasGluyas Early WW15
Gluclub Aroona
Clubhead Raven Tadorna 3* Cascades
Tincurrin
4*
Chile 1B Datatine Inia 66
3Ag3-Condor
Insignia Halberd 3*
Falcon

Figure 4 Pedigrees of recent prominent varieties of wheat grown in Eastern and Western Australia. Pedigrees are read from left to right. For example,
Raven was crossed with the line WW15 to produce the variety Aroona. The line WW15 was crossed with WW80 and then crossed again (double
backcross, 2*) to produce the variety Condor. The variety Meering was selected from Condor. Condor was a parent entering into the pedigrees of Banks
and Cook. A sister line of Condor was a parent that also entered into the pedigrees of Banks and Cook.
 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries 415

allowing the beds to dry prior to mechanical header harvesting.

Yields are high by world standards, averaging over 9 tonnes per
hectare (Table 5), with some growers achieving 13 tonnes per
hectare. Rice is generally delivered at relatively high moisture
(2022%) to avoid cracking of the kernels. Aerated storage is
used to further reduce the moisture to about 15% prior to milling.
All deliveries are sampled on delivery and are analyzed for mois-
ture and protein contents by near-infrared transmission spectros-
copy. Samples are also checked for variety and for defects.
Most of the rice grown in Australia is from medium- and
long-grain Indica varieties. The long-grain varieties include the
fragrant types. The hulls that are removed in the milling process
are converted by incineration to high-carbon ash, which is used
by the steel industry in Australia and overseas. The ash is used to
coat the surface of open-hearth furnaces, thus trapping the heat
and improving fuel efficiency. Fireproof building materials
have also been developed using this ash as a starting material.
Rice accounts for about 8000 tonnes of the 9000 tonnes of
Figure 5 A quarter-scale model of the small-scale flour mill used by cereal-grain production in Fiji (Table 1).
Guthrie to evaluate the milling quality of the small amounts of grain
coming from Farrers wheat breeding program. It is now on show beside
the commercial-size roller Ganz mill in the Foundry Museum in Budapest.
Coarse Grains
Table 4 Production, in kilotonnes, of major cereal crops in Australia Barley
for successive crop years
Barley is second in importance to wheat, based on volume and
Crop 2010 2011 2012 quality (Tables 2, 4, and 5). Like wheat, yields of barley are
moderate by world standards (Table 5). Export markets are
Wheat 22 138 27 410 29 905 mainly China, the Middle East, and Japan. Australia is the
Barley 7294 7995 8221
second most important country in world malting-barley
Oats 1374 1128 1262
trade, depending on seasonal fluctuations. The premium use
Maize 328 357 451
Triticale 502 284 285 of barley is for malting to produce beer. In addition to the
Sorghum 1598 1935 2239 export of malting barley, malt produced in Australia is
Rice, paddy 197 723 919 exported for beer production overseas. Varieties are bred spe-
cifically for the purpose of malting, based especially on the
Source: http://faostat.fao.org. production of brewery extract for fermentation purposes after

Prime Hard and Hard

Hard and Fair Average Quality
Fair Average Quality some Hard
Fair Average Quality
Soft and Fair Average Quality
Export terminal.

Figure 6 Types of wheat quality according to the fair average quality system of the mid-twentieth century in Australia. Map provided by John Moss.
416 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries

Table 5 Mean yields of crops in tonnes per hectare for the 2012 crop
year

Crop Australia New Zealand World

Wheat 2.1 8.9 3.1

Barley 2.2 6.7 2.7
Rapeseed 1.5 1.1 1.9
Sorghum 3.4 1.5
Seed cotton 4.8 2.2
Rice (paddy) 8.9 4.4
Maize 6.5 10.9 4.9
Oats 1.7 4.7 2.2
Peas (dry) 1.4 3.1 1.5

Source: http://faostat.fao.org.

Figure 7 Arabic pocket breads baked experimentally by an expert from

the Middle East in the laboratories of the Bread Research Institute of
Australia in the 1970s. This type of flat bread comes from the oven puffed
into a sphere. On cooling, it returns to its flat-bread form. Advice from
such experts has assisted Australia in recent decades in providing wheat
of the quality suited to specific market requirements.
Figure 9 The range of food products made from Australian wheat. In
addition to the many forms of leavened bread, Australian export wheat
is used for products such as Chinese steamed bread (white sphere
at lower left), Arabic flat breads (lower left), noodles (at right), crisp
breads (lower center), and pasta (lower right).

malting. These are all two-row types; Australian six-row barleys

Figure 8 The grain-storage facility in Perth, Western Australia. The very
large storage capacity of this recent construction was warranted by
the recent increase in grain production in Western Australia. The many
storage cells are designed to accommodate the diversity of grain species
and grades.
are of feed quality.
Animal feed is the destination of barley that does not meet
the stringent requirements of the malting grades, due to fail-
ures in meeting specifications for variety, protein content, or
absence of defects. Feed barley, together with sorghum and
downgraded wheat are the main grains for the animal-feed
industry, their importance being greater because of the lower
production of maize in Australia.
Barley is also used for human food purposes both domes-
tically and in export markets. Japan has emerged as a major
user of food-grade barley. Much of the food use involves pear-
ling, to remove the bran and husk layers. In other cases, flour is
produced and some of this is becoming used to produce high-
fiber pasta products, taking advantage on barleys high-fiber
content in its cell walls (b-glucans and pentosans). The pro-
duction of distilled beverages is another significant use of food-
grade barley.
 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries
Oats
Most of the oat crop (Tables 2, 4, and 5) is used within
Australia. It may serve on-farm as a forage crop to be grazed
when plants are young, with the option of allowing the plants
to mature and produce harvestable grain. Some varieties are
suited to milling and processing for food purposes, but much
of the oat crop is used as animal feed in its various forms
forage, hay, and grain.
Sorghum
Sorghum is a feed grain. Much of the crop is exported for this
purpose. The main customers are Japan and Taiwan. Produc-
tion is mainly confined to the warmer subtropical parts of
Australia.
Cereal Rye
Flour produced from rye grain is used in Australia for
European-style breads, biscuits, crispbreads, multigrain break-
fast cereals, muesli bars, pet food, and special dietary foods that
provide higher lysine, pentosan, and b-glucan content than
other cereals. Production is minor in volume.
Triticale
The wheatrye hybrid triticale has enjoyed reasonable popu-
larity in Australia. Annual production varies, recently peaking
in 2010 at about half a million tonnes (Table 4), making
Australia the worlds fourth largest producer of triticale,
depending on seasonal fluctuations. Triticales attraction has
mainly been as an alternative to wheat for the stockfeed indus-
try, which is the major domestic end user. However, efforts
have been made to develop niche markets for human food.
Maize
Maize is a relatively minor crop in Australia, with annual
production ranging up to half a million tonnes (Table 4),
less than the production figures for the major cereal grains.
However, grain yields are good. About 10% of the maize crop
is exported. About one-quarter of the crop is used domestically
as human food, mainly for breakfast cereals, sweet corn/vege-
table uses, corn-based foods, and industrial manufacture of
corn starch. The remainder is used domestically as stockfeed.
Industrial use of maize is minor compared to northern-
hemisphere maize use, because Australian starch production
is mainly based on wheat. Maize processing into starch relates
mainly to special products such as high-amylose starch.
Pulses
Pulse production in Australia has expanded greatly since 1980
in response to changing farming systems and increasing
world demand for vegetable protein. The main species are
lupin, chickpea, lentil, field pea, and faba bean (Table 2).
Lupin and field pea have been grown in Australia since the
nineteenth century when they were used mostly as animal
feed. There was little plant breeding effort at this time: one
417
of the pea varieties grown in the early twentieth century, Dun,
was still being grown in the 1980s. This began to change with
the work of John Gladstones on lupin at the University of
Western Australia and later in the WA Department of Agricul-
ture beginning in the 1950s, in which he took what was
essentially a wild plant, Lupinus angustifolius, and by gradually
incorporating genes for low seed alkaloids, soft seed, non-
shattering pods, white flowers, and early flowering, created
the modern crop-type narrow-leafed, or Australian Sweet,
lupin beginning with the release of Uniwhite in 1967. Field
pea breeding resumed in Australia in the 1970s, and chickpea
and lentil breeding began in the 1980s. All of these pulses are
susceptible to devastating diseases, and improving disease
resistance has been an important objective of these breeding
programs, as well as improving adaptation to Australian
conditions.
Pulses in Australia are grown in rotation with cereals and
improve production by providing fixed N and reducing cereal
disease levels. Production expanded rapidly in the 1980s and
1990s as cereal production intensified, whereas legume-based
pastures, which had formerly played a similar rotational role,
declined in response to the changing economic environment
of Australian farming.
There are geographic differences in pulse production
across the country. Narrow-leafed lupin is the predominant
pulse in Western Australia; white or albus lupin (L. albus) is
grown mainly in southern New South Wales, desi chickpea in
northern New South Wales, and the other species in South
Australia, Victoria, and southern New South Wales. Produc-
tion of lupin, field pea, and lentil declined between 2000 and
2012 in response to changing grain prices relative to wheat
and canola, which has displaced pulses to some extent.
Increasing weed control costs as herbicide-resistant weeds
become more prevalent has contributed to declining lupin
production in Western Australia, and more frequent drought
during the past decade has also been important. On the other
hand, production of chickpea has more than quadrupled in
the same period.
A high proportion of Australian pulse production is
exported. Field pea, lentil, and chickpea mostly go to the
Indian subcontinent where they are used in human food,
although a significant proportion of field pea is used domesti-
cally in pig and poultry rations, especially in Eastern Australia.
Australia produces more than 70% of the worlds lupin and
about 50% of this is exported. Most goes to Europe or northern
Asia where it is used for animal feed. Australian lupin is also
used for aquaculture feed, and there has been considerable
effort over the past decade to develop human food uses for
lupin, including fiber and protein additives, which can be used
in a wide range of manufactured foods. However, these are
currently only minor uses.
Oilseeds
The canola (rapeseed) crop has expanded dramatically since
the 1990s, production having tripled in the period 199699. It
was about 2.4 Mt in 1999, rather less than the figure given in
Table 2 for 2012, and is now the third-largest crop in Australia
after wheat and barley. It performs a similar role in crop
418 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries
rotations to pulses (although it does not fix atmospheric N),
and its expansion has to some extent been responsible for the
stagnation of pulse production in the past decade. Canola is
one of the two crops in which genetically modified (GM)
varieties are approved for commercial production in Australia
(the other is cotton). Currently, between 10% and 20% of
production is from varieties with the Roundup Ready trait.
In recent years, about two-thirds of Australian production has
been exported, and Australia is the worlds second-largest
exporter of canola after Canada. Cottonseed production is the
volume of white seed, after allowing for delinting and de-
hulling. About a quarter of this is crushed domestically for oil
production. Yields are good (Table 5).
Production volumes are modest for sunflower and soybean
(Table 2). Peanut production is well below these at about
25 000 tonnes. Lesser oilseeds include linseed and safflower.
Australia is a significant exporter of oil crushed from these
various oilseeds, especially cottonseed. The residues from oil-
seed crushing make important contributions to the stockfeed
industry.
Consumption
Australians consume about 70 kg of wheat flour per person
per year, mainly in the form of baked goods, but flour also
goes into many processed foods. A significant part of this
consumption (based on statistics) is due to the importance
of the starchgluten industry, which processes dough to
separate the starch fraction, using the A starch for paper
manufacture. Wheat starch is also a significant food ingre-
dient. The remaining gluten is dried and used as an additive
for bread manufacture (both domestically and as a signifi-
cant export commodity) and also for incorporation in a
range of foods. The residue of starch is fermented to etha-
nol for blending with petrol for vehicle fuel. Another sig-
nificant use of grains in Australia is the breakfast-cereal
market, which is valued at $800 million. These and other
grain-based foods represent an important part of Australias
exports of manufactured foods, primarily to the Pacific
Islands and to Asia.
New Zealand
Significant volumes of wheat, barley, and maize are produced
in New Zealand, as is shown in Table 2. Grain yields are
very high by world standards (Table 5). Deregulation of the
wheat industry some years ago accounted for significant
changes in the structure of the industry, increasing competition
and the likelihood of importation of grain. In contrast to
Australia, adequate on-farm storage for the crop is common
(Figure 10), so that the grain buying and transport systems are
different from those in Australia. Most of the New Zealand
harvest is later than in Australia, where early deliveries may
start in October. On the other hand, harvest time in the south
of the New Zealand South Island may be as late as Easter time
(late in the southern-hemisphere autumn).
Figure 10 Wheat field and on-farm storage in the south of the South
Island of New Zealand. As late as April (autumn), the standing grain is still
waiting to be harvested.
Future Prospects
The future for international grain trading is difficult to predict,
largely due to the entry of new exporting nations and the
deregulation of marketing strategies in several other countries.
Looking further ahead, climate-challenge issues will affect pro-
duction and quality characteristics. Likely climate changes
include more frequent heat episodes and rainfall that is vari-
able and less reliable. One of the results may be the need to
move away from traditional growing regions for some grain
species. Rising sea levels may reduce the amounts of arable
land in Pacific Islands nations.
It is inevitable that there will be a continuing rise in the
level of carbon dioxide in the Earths atmosphere, although the
causes and the rate of increase are subject to discussion at
present. Many studies have already shown that the resulting
carbon fertilizer effect produces increased yields of grain and
biomass, accompanied by lower protein content for the result-
ing grain. For grains like wheat, a significant reduction in
protein content may require changes in baking procedures,
such as the use of added gluten to bolster dough properties
or use of the Chorleywood bread process, which was devel-
oped originally to accommodate low-protein wheat in the
United Kingdom. Wheat breeders may also need to incorporate
gluten genes associated with greater dough strength.
The many experimental trials of elevated CO2 have pro-
vided a basis for modeling to extrapolate predictions of future
effects to a wider range of environments. For example,
Figure 11 illustrates expectations for the state of Victoria in
southeastern Australia. It takes into account the rising levels of
CO2 together with other expectations, such as heat episodes,
rainfall, and known soil types. Although large areas are
expected to enjoy higher crop yields in 70 years time, there
are significant areas where that result may not be expected.
Research and development is ongoing, especially in
Australia and New Zealand, with outcomes expected to benefit
both grain yield and quality. New quality types include the
production of low-glycemic-index wheats (based on high-
amylose wheats) that will address chronic health issues such
as diabetes, obesity, and cardiovascular disease. Research on
 GRAINS AROUND THE WORLD | Grain Production and Consumption in Oceania: Australia and Pacific Countries
N
0 98
kilometers
Figure 11 Predicted changes in crop yields for the Australian state of Victoria, based on modeling of rising levels of CO2 together with other
expectations, such as heat episodes, rainfall, and known soil types. Illustration, courtesy of G. OLeary and coworkers.
canola is providing lines capable of producing omega-3 fatty
acids. The mining of novel genes is being facilitated by the
establishing the Australian Grains Genebank in Horsham, Vic-
toria. It houses about 300 million seeds from around the
world, for use in current and future plant breeding programs.
Exercises for Revision
What features distinguish grain production, quality, and
consumption in your region from these factors in Oceania
(or choose one of the Oceanian countries).
What advantages are claimed for Australian export wheat?
What are the reasons for determining the variety identity in
the Australian wheat harvest?
Name some of the improvements made to Australian
wheats by breeding and selection.
What uses of barley and wheat are described?
Exercises for Readers to Explore the Topic Further
What factors could account for the differences in grain
yields for the various grain species in Table 5, contrasting
Australia, New Zealand, and world averages?
Find out about the development and expansion of wheat in
your region. How did their early times contrast with the
origins of the Australian wheat industry?
How would starch from wheat (as is usual in Australia)
differ from the commercial production of starch from
maize in many northern-hemisphere countries in produc-
tion methods and in quality?
Find out about the relative advantages of the oilseeds men-
tioned with respect to the amount and quality of the oils
produced.
How does the value of the respective residues after oil
removal affect the overall economics of oilseed utilization?
See also: Grain Harvest, Storage and Transport: Barley:
Harvesting, Storage, and Transport; Grain Marketing and Grading:
419
Crop yield still increasing by year 2070
Decreasing crop yield by year 2070
Decreasing crop yield by year 2050
Decreasing crop yield by year 2030
196 Decreasing crop yield by year 2015
Identification of Varieties of Food Grains; Wheat: Grading and
Segregation; Wheat: Marketing; Non-food Products from Grains:
Fuel Alcohol Production; The Cereal Grains: An Overview of the
Family of Cereal Grains Prominent in World Agriculture; Barley: An
Overview of a Versatile Cereal Grain with Many Food and Feed Uses;
Wheat: An Overview of the Grain That Provides Our Daily Bread; The
Oilseeds: Canola: Overview; Cottonseed: Overview; Wheat
Processing: Gluten and Modified Gluten.
Further Reading
Anonymous (1994) Australian Grains. Melbourne, VIC: Morescope Publishing.
OBrien L and Blakeney AB (eds.) (2000) An Introduction to the Australian Grains
Industry. Melbourne, VIC: Royal Australian Chemical Institute.
OBrien L, Morell M, Wrigley C, and Appels R (2001) Genetic pool of Australian wheats.
In: Bonjean AP and Angus WJ (eds.) The World Wheat Book, pp. 603640. Paris:
Lavoisier Publishing.
OLeary G, Christy B, Weeks A, et al. (2011) Downscaling global climatic predictions
to the regional level: A case study of regional effects of climate change on
wheat crop production in Victoria, Australia. In: Shyam S, Yadav RJR, Hatfield JL,
Lotze-Campen H, and Hall AE (eds.) Crop Adaptation to Climate Change,
pp. 1226. Chichester: Wiley.
Quail KJ (1996) Arabic Bread Production. St. Paul, MN: American Association of Cereal
Chemists.
Wrigley CW (1978) W.J. Farrer and F.B. Guthrie: The unique breeder-chemist
combination that pioneered quality wheats for Australia. Records of the Australian
Academy of Science 4: 725.
Wrigley CW (1981) Farrer, William James. In: Nairn NB and Serle AG (eds.) Australian
Dictionary of Biography: 18911939, pp. 471473. Melbourne: Melbourne
University Press.
Wrigley CW (1983) Guthrie, Frederick Bickell. In: Nairn NB and Serle AG (eds.)
Australian Dictionary of Biography: 18911939, vol. 9, pp. 143144. Melbourne:
Melbourne University Press.
Wrigley CW (2005) Contributions by Australians to wheat quality research. In: OBrien L
and Blakeney AB (eds.) An Introduction to the Australian Grains Industry,
pp. 3341. Melbourne: Royal Australian Chemical Institute.
Wrigley CW and Rathjen A (1981) Wheat breeding in Australia. In: Carr DJ and
Carr SGM (eds.) Plants and Man in Australia, pp. 96135. New York: Academic
Press.
Wrigley CW, Tomoskozi S, and Bekes F (2011) Hungarian-Australian
collaborations in flour milling and test milling over 120 years. Cereal
Research Communications 39(2): 216225. http://dx.doi.org/10.1556/
CRC.39.2011.2.5.
 Grain Production and Consumption: South America
OR Larroque, CSIRO Agriculture Flagship, Canberra, Australia
JC Catullo, INTA, Buenos Aires, Argentina
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Brazil and Argentina are the major producers and traders in
the region.
Farmland will continue to increase as more virgin land is
transformed for agriculture use.
GMOs are widely used in South America.
Technological advances in mechanization (i.e., no-tillage)
proved crucial to farming success.
Soybean will continue to be the major cash crop in the
region.
Constant monitoring to assure sustainability is needed.
Cereals are an important part of the South American diet.
Breakfast cereals are not widely consumed.
Pseudocereals like quinoa and amaranth are becoming
more popular.
Learning Objectives
Current status of South American production, trading, and
consumption of grains.
Prologue
Since the onset of the 2008 global economic crisis, investors
favored more secure alternatives than the crumbling financial
sector options. As a result of that and on the back of increasing
international food prices, agriculture became an attractive
investment option worldwide and particularly in South
America.
South America is one of the regions with greater aptitude
for providing food and renewable energy to the rest of the
world. The whole region possesses plenty of arable land and
renewable water resources. Increases in harvested area
(Table 1) and improvements in average yields of the major
crops (Table 2) have boosted the regions production (Table 3)
in the last decades. This was possible thanks to better seed
varieties, mechanization, no-tillage farming, and widespread
and more efficient agrochemicals and fertilizer usage.
Fifty-one percent of the worlds soybean production and
11% of maize are harvested in South America (average
201012) per year. Wheat accounts for just over 5% of the
world production. While maize and wheat increased their
share in the last decade by 1.3% each, soybeans share of the
global market production increased by a whopping 7%.
The agricultural boom has prompted the governments of
the region to adapt their policies to the emerging situation in
grain trading. For example, Argentina implemented measures
since 2007 to unpeg high global food prices from local prices.
The country increased soybean export taxes and limited exports
420 Encyclopedia of Food Grains, Second Edition
(namely, wheat and maize) in order to secure a well-supplied
local market.
Cereals share in total food supply varies in the region from
29.0% in Argentina to 48.3% in Guyana. Argentina, Bolivia,
Brazil, Chile, Peru, and Uruguay have wheat as the main staple
cereal component of their diets, while maize is the prevalent
cereal component in Colombia, Paraguay, and Venezuela. Rice
is the more ubiquitous cereal component of the diet in Ecua-
dor, Guyana, and Suriname (Figure 1).
Argentina
The Argentine Republic is the second grain producer and the
main grain exporter of South America (Figure 2). Technolog-
ical advances, better seed quality, genetic improvement, and
better crop management were all important factors in the
resurgence of agriculture as one of the main driving forces of
the countrys economy.
From a total continental area of 273 670 000 hectares,
Argentina dedicates 38 000 000 hectares to agriculture (2011).
Ten years ago, the arable land was 27 000 000 hectares. In
contrast, rural population continues to decrease (10.73% of
total population in 1996; 7.27% in 2012).
The share of agriculture value added in total GDP is gradu-
ally returning to increase (from a lowest value of 4.79% in
1999) although not to the levels of 20% that they had up to
half a century ago. Agriculture value added (% of GDP) in
Argentina was 10.61 as of 2011.
Wheat, maize, sunflower, and soybean account for 90% of
all the grain produced in the country, making the top ten world
producers and the top five exporters list for each of the crops.
Other important grain crops are sorghum (5 200 000 tonnes
in 2012) and barley (5 500 000 tonnes in 2012). Argentina
ranks fifth in the world, with more than 8% share of sorghums
global production, and is the second largest exporter behind the
United States. Chile, Japan, Colombia, Peru, Germany, and
Ecuador are the main buyers of Argentine sorghum.
In the last 15 years, there have been steady increases in
exports of most of the grains, particularly coarse grain and oil
crops.
The main production area is the Pampas, once called the
Granary of the World, a rich humid plain in the central eastern
part of the country, which concentrates 80% of grain production.
Wheat is the top overall commodity available for consump-
tion (2009) with 678 kcal per capita per day. Maize and rice are
the joint second grain and sixth overall (81 kcal per capita per
day). Cereals account for 29.0% of the total food supply
(2009). Total wheat flour consumption is 94 kg per capita
from which 64 kg corresponds to continental bread, 10 kg to
pasta, 7 kg to crackers, and 4 kg to sliced bread, according to
statistics from the Milling Industry Federation (FAIM).
http://dx.doi.org/10.1016/B978-0-12-394437-5.00050-4
 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America 421

Table 1 Grain-cultivated area in South American countries

Beans, Rice, Sunflower

Country Barley dry Maize Oats Quinoa paddy Sorghum Soybean seed Triticale Wheat

Argentina 1121 274 3384 227 236 971 18 749 1687 4189
Bolivia, Plurinational 53 52 331 6 64 178 105 1012 182 175
State of
Brazil 92 3275 13 375 175 2615 703 24 078 72 45 2071
Chile 17 11 127 97 24 3 23 260
Colombia 3 116 510 3 488 13 33 6
Ecuador 25 46 370 1 1 365 9 44 8
French Guiana 3
Guyana 3 132
Paraguay 61 812 70 24 2826 104 572
Peru 151 81 493 11 2 36 380 1 151
Suriname 55
Uruguay 64 5 105 23 180 32 952 5 373
Venezuela, Bolivarian 55 664 245 243 23 62 1
Republic of
World 48 401 30 040 170 865 9462 101 162 759 40 410 104 283 24 681 3842 218 197

Average 201012 (thousands of hectares).

Source: FAOSTAT.

Table 2 Yield of major producers of grains in South America

Country/yield 2003 and 2012 Maize 03 Maize 12 Wheat 03 Wheat 12 Rice 03 Rice 12 Soybean 03 Soybean 12

Argentina 6.48 7.34 2.54 2.97 2.80 2.66

Bolivia, Plurinational State of 2.32 2.20
Brazil 3.73 5.01 2.40 2.32 3.25 4.81 2.80 2.63
Chile 10.00 10.72 4.32 4.95
Colombia 6.79 7.72
Paraguay 2.38 3.90 2.20 2.46 2.85 2.78
Peru 5.86 4.14
Uruguay 2.77 2.85 2.55 2.65
Venezuela, Bolivarian Republic of 3.43 4.29
World 4.46 4.94 2.70 3.12 3.95 4.39 2.28 2.37

Years 2003 and 2012 (tonnes per hectare).

Source: FAOSTAT.

Table 3 Grain production in South American countries

Country/production Beans, Rice, Sunflower

avg 201012 Barley dry Maize Oats Quinoa paddy Sorghum Soybeans seed Triticale Wheat

Argentina 4180 340 24 059 492 1520 4429 51 019 3078 13 656
Bolivia, Plurinational 49 61 915 6 37 454 401 2206 238 235
State of
Brazil 280 3 60 774 387 12 035 1834 69 757 95 110 5414
Chile 98 20 1430 465 125 6 108 1438
Colombia 5 133 1643 4 2001 59 72 11
Ecuador 18 13 1004 1 1 1583 13 72 7
French Guiana 10
Guyana 5 582
Paraguay 53 3178 368 148 8040 165 1421
Peru 210 91 1578 12 42 2825 3 220
Suriname 227
Uruguay 210 3 455 47 1405 122 2119 6 1167
Venezuela, Bolivarian 47 2421 1334 481 38 54
Republic of
World (total) 129 693 23 173 870 060 21 114 80 713 984 58 578 260 075 36 418 13 652 453 350

Average 201012 (thousands of tonnes).

Source: FAOSTAT.
422 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America
100
90
80
70
60
% 50
40
30
20
10
0
Argentina Bolivia Brazil Chile Colombia Ecuador Guyana Paraguay Peru Suriname Uruguay Venezuela
Figure 1 Share of main cereals in the South American diet. Source: FAOSTAT (2009).
According to the same source, the country has 33 000 bakeries
where artisan continental bread is baked daily. Argentina is
currently the fifth largest world exporter of wheat flour.
Argentinas undernourishment figures are not significant,
comprising <5% of the population in 201012.
Bolivia, Plurinational State of
Landlocked Bolivia is the least developed country of the region,
with a large proportion of its population living in rural areas
(32.54% in 2012). Arable land has increased in recent years
from 2.59 million hectares in 1996 to 3.84 million hectares in
2011. This increase was mainly made at the expense of forest
cover. Agriculture employs 40%of the workforce (down from
45% in 1997). While the GDP has doubled between 2005 and
2010, the share of agriculture value added in total GDP has
decreased, continuing a trend that saw its figures going down
from nearly 20% in 1995 to just above 10% in 2010.
In recent decades, this Andean country has consolidated a
strong growth in oilseed crop production, particularly soybean
with planted areas exceeding 1 000 000 hectares and total pro-
duction of 2 298 857 tonnes in 201112. Soybean is the largest
agricultural export and the third largest Bolivian export.
Santa Cruz de la Sierra is the main producing area with an
average production yield of 16002400 k ha1. There are two
crops per year: the most important in summer (November/
December) and also in winter (June/July).
The second most important grain crop is maize
(1 006 000 tonnes in 2012) followed by sorghum
(478 000 tonnes) and paddy rice (440 000 tonnes).
Wheat
Rice (milled)
Maize
South
America +
(Total)
Wheat is the top commodity available for consumption
(2009) with 406 kcal per capita per day. Maize is the second
(302 kcal per capita per day) and rice third (280 kcal per capita
per day). The prevalence of undernutrition is very high at 24%
(201012).
Dry bean production in 2012 was 60 000 tonnes.
Bolivia is also the largest producer and exporter of quinoa, a
high-protein grain native to South America that was the staple
food of the Incas. The UN declared 2013 the Year of Quinoa.
Unfortunately, higher international prices for the crop are
leaving the poorest segments of the population unable to
purchase quinoa anymore.
Brazil
The South American giant is the largest economy of the region
and the major grain producer. Brazilian agriculture has chan-
ged dramatically in the last 30 years. From being a major food
importer (Figure 3), the country also became a top grain
exporter (Figure 1). Brazil is the first of the big exporters to
be in a tropical area and not a temperate area (like it is the case
with the United States, Canada, Australia, Argentina, and EU).
The arable land has increased from 57 860 000 ha in 1996
to over 70 000 000 ha in 2011. It corresponds to 8.3% of the
total country area. Furthermore, it has over 400 000 000 ha of
potentially arable land, the largest area for any country in the
world according to FAO. Most of this land is in the Brazilian
savannah (the Cerrado).
 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America 423

35 000
32 986

30 000

26 832

25 000

20 000

15 806
Thousands of tonnes
15 000 14 355

10 820
10 000 9487
8411

5010
5000 4417

0
Soybean Soybean cake Maize Soybean cake Soybean Maize Wheat Soybean Soybean oil
Brazil Argentina Argentina Brazil Argentina Brazil Argentina Paraguay Argentina

Figure 2 Major grain commodities exports South America. Source: FAOSTAT (2011).

The most recent data show that primary agriculture
accounted for 5% of Brazils GDP and agricultural population
nears 20 000 000 people (12% of total population).
Brazil is second largest producer of soybean oilseed
(88 000 000 tonnes) and the third largest producer of maize
(70 000 000 tonnes) in the world, behind the United States
and China. In 2011, maize exports accounted for
9 486 914 tonnes (second in the region and third in the
world), and soybean exports were just below
33 000 000 tonnes (number one exporter in the world).
The country is the sixth largest importer of wheat
(5 740 753 tonnes in 2011) although it also exports
(2 350 000 tonnes in 2011). Brazil is the third largest producer
of pasta in the world.
Rice is a staple food in the country (352 kcal per capita per
day), so its consumption is correlated with the population
growth. Most of Brazilian rice is cropped in uplands, particu-
larly in the northern and midwestern regions. Production of
rice (milled) was 8 200 000 tonnes in 2011. Brazil is the ninth
world producer.
Other grain commodities available for consumption are
wheat (384 kcal per capita per day), soybean oil (336 kcal per
capita per day), maize (209 kcal per capita per day), and beans
(151 kcal per capita per day). Brazils traditional, basic meal
consists of rice and beans (feijoada). Brazil is the second
world producer of dry beans, behind India. Production figures
for 2012 were nearly 3 000 000 tonnes. Consumption of
16.30 kg per capita per year was recorded for 2009.
Although the importance of breakfast cereals in the region
is limited, Brazil and Venezuela show some increase in con-
sumption. From 2011 to 2016, it is predicted that Brazils
market will increase by 27%. Increase in consumption is asso-
ciated with upper and middle-upper income segments of the
population. On the other end of the spectrum, breakfast cereals
are still very expensive for lower-income earners when com-
pared with traditional breakfast meals.
The prevalence of undernutrition has dropped from 12% in
19992001 to 7% in 201012. The Zero Hunger Program
launch in 2001 by the then-presidential candidate Lula da
Silva has successfully helped in the improvement of these
figures. Family farming, the main pillar of food security in
the country, is responsible for the domestic production of
70% of beans, 46% of maize, 34% of rice, and 21% of wheat.
Brazil legislation now permits the use of GMOs. Since the
full lifting of the ban in 2005, Brazil has turned into the second
largest producer of GM crops in the world, behind the United
424 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America

7000

6000 5740

5000

4000

Thousands of tonnes
3000 2820

1908
2000
1684
1535
1299 1215
1014 1004
1000

0
Wheat Maize Maize Wheat Wheat Maize Wheat Soybean cake Soybean cake
Brazil Colombia Peru Peru Colombia Venezuela Venezuela Colombia Peru

Figure 3 Major grain commodity imports into South American countries. Source: FAOSTAT (2011).

States. Over 80% of the soybean crop is GM, and they are
planning to bring to the market a GM bean with resistance to
the golden mosaic virus.
Chile
Located on the western coast of the continent, facing the
Pacific, the Republic of Chile occupies a narrow strip of land
between the sea and the Andes mountains.
The share of agriculture value added in total GDP is very
little (<5%) and the rural population is just above the 10%
mark. The arable land has diminished in the last years, from
2 000 000 hectares in 1996 to 1 320 000 hectares in 2011.
The main grain crop is wheat (1 575 822 tonnes in 2011).
A quarter of this production comes from 40 000 small farmers
(subsistence group, no or little alternative crop to grow).
A little over 550 farmers produce 50% of the wheat and around
5000 farmers are considered medium size producers. Maize
(1 493 292 tonnes in 2012) is the second crop in importance.
Chile is the top producer of oats (450 798 tonnes in 2012) in
the region and within the top five in the world.
To fulfill internal demand, wheat (655 527 tonnes), maize
(668 151 tonnes), and sorghum (636 112) are three of the
most imported commodities in the country (2011).
Wheat is the top overall commodity available for consump-
tion (2009) with 850 kcal per capita per day. Maize is the
second grain and third overall (211 kcal per capita per day)
and rice the third and eighth, respectively (90 kcal per capita
per day). Cereals account for 42.3% of the total food supply
(2009). Chileans consume an average of 86 k of bread per
capita per year (200110), making them the largest consumers
of bread in the region and second in the world, behind
Germany. The most consumed types are the soft bread marra-
quetas and the flat round hallullas. Chile fortifies basic food
(in this case, wheat flour) with micronutrients (iron).
Chiles undernourishment figures are not significant, com-
prising <5% of the population in 201012. In 199092, 8.1%
of the population was undernourished.
Colombia
Situated in the northwestern part of South America, Colombia
is the fourth largest country of the region, after Brazil,
Argentina, and Peru. The arable land fluctuates around 2% of
the total area of the country. The eastern plains region called
the Altillanura has potential for future agricultural expansion.
The share of agriculture value added in total GDP has
abruptly dropped from the turn of the millennium. In 2011,
 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America
Colombias agriculture contributed nearly 7% of total GDP.
Nevertheless, agriculture continues to be an important source
of employment (24% of the economically active population).
The most important grain crop is rice (paddy
2 000 000 tonnes; milled 1 320 000 tonnes in 2011), which
ranks sixth among the top ten commodities produced in
Colombia. Most of the crop is grown under irrigation. The
harvested area averages just over 400 000 ha in recent years.
Maize contributed with a production of 1 700 000 in 2011.
The harvested area remains steady at 500 000 ha in recent
years. Around 10% of the planted area uses biotech seed and
is expected to increase in the future.
Colombia is the second largest importer of the region,
behind Brazil. Maize imports have been well above
3 000 000 tonnes in recent years (3 900 000 tonnes in 2013)
positioning the country as the largest maize importer in
South America. Ninety-five percent of maize imports (mainly
yellow) are used in the feed industry.
Soybean is being used in rotations with maize, but produc-
tion is not significant (86 634 tonnes in 2012)
Wheat imports average 1 500 000 tonnes per year. Bread
production demands 75% of the wheat, while the pasta indus-
try requires 15% and the cookies and pastry industry the
remaining 10%.
Rice (358 kcal per capita per day), maize (333 kcal per
capita per day), and wheat (222 kcal per capita per day) are
the main commodities available for consumption. Colombia is
the third largest producer of beans in South America
(133 000 tonnes in 2012) and ranks fourth in consumption
with 3.5 kg per capita per annum.
Prevalence of undernutrition (201012) remains steady
at 13%.
Ecuador
The Republic of Ecuador has borders with Colombia on the
north, Peru on the east and south, and the Pacific Ocean on the
west. Arable land accounted for 1 160 000 hectares in 2011
(4.7% of the total area). Agricultural population is still high
at almost 20% of the total population, but the numbers have
decreased quite constantly at a rate of 1% of the population
size per year.
The share of agriculture value added in total GDP is 10%.
Rice (paddy, 1 655 535 tonnes in 2012) and maize
(864 445 tonnes in 2011) are the main crops grown in Ecua-
dor. This satisfies internal rice demand (which is a very sensi-
tive issue in Ecuador), but not maize, which is also imported to
fulfill demand (543 817 tonnes in 2011). Imported maize is
used mainly by the feed meal compound industry and the
related poultry, egg, and pork industry. Recent crop forecasts
predict large increases in maize harvest (over 1 200 000 tonnes)
and subsequently less need for imports.
Ecuador imports most of its wheat (565 523 tonnes in
2011). Despite government measures tending to boost wheat
production, domestic-grown areas are still insignificant. The
importation of soybean cake amounted for just over
600 000 tonnes in 2011 (the fourth importer in the region,
after Peru, Venezuela, and Colombia). Sorghum imports
amounted for 26 264 tonnes in 2011.
425
Dry beans production averaged 13 000 tonnes for 201012.
The main grain commodities available for consumption in
2009 were rice (milled) at 431 kcal per capita per day and
wheat (295 kcal per capita per day). Soybean oil availability
for consumption was 120 kcal per capita per year. Cereals
share was 34.4% in total food supply (2009).
The prevalence of undernutrition was one of the highest in
the region with 18% for the period 201012, slightly down
from an average of 21% (19992009).
French Guyana
Agriculture is concentrated in coastal areas (10 000 hectares of
arable land). Rural population accounts for 23% of the total
population.
The main and almost only grain crop grown in the country
is rice, with a production of 9587 tonnes in 2011 (130th in the
world). Production has dramatically decreased in the last
decade (31 000 tonnes in 2001).
Guyana
Annual crops are grown on the narrow coastal strip. Arable
land represents 2% of the total country area. Rice is almost the
only grain produced, with nearly 600 000 tonnes harvested in
2011. There are two crops per year.
Agricultural value added in total GDP decreased from
around 40% in 1995 to just over 20% in 2010.
The main grain commodity available for consumption in
2009 was rice (796 kcal per capita per day) followed by wheat
(483 kcal per capita per day). Cereals account for 48.3% of the
total food supply (2009), the highest in the region.
Paraguay
Paraguay is landlocked between the two South American pow-
erhouses: Brazil and Argentina.
Agriculture, including subsistence farming, is the main eco-
nomic activity. From a population of nearly 7 million in 2012,
agricultural population represented 28% of the total and 24%
of the total labor force. The share of agriculture value added in
total GDP was 23.1% as of 2011, positioning the country in the
first place for this indicator in the region. Gross domestic
product and gross national income per capita have both dra-
matically increased since 2005. In 2013, the Paraguayan
economy is expected to have the largest GDP growth (13%)
of the region.
Arable land has increased mainly at the expense of forest
cover and permanent crops. Like in other countries in the
region, this was mainly due to the economic importance of
cash crops. The eastern region is the major production area in
the country, with a mixture of traditional small farms and well-
equipped medium/big farms. Main producing states are Alto
Paranaa, Itapua, and Canindeyu.
Soybean is the main agricultural commodity in both
production (8 309 793 tonnes in 2011) and value. Constant
increases in planted areas and excellent yields have skyrocketed
426 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America

this crop in detriment of maize (third place, 3 345 877 tonnes),
wheat (fifth place, 1 461 020 tonnes), rice, and sorghum. Para-
guay is the fourth most important exporter of soybeans in the
world.
Maize is the top commodity available for consumption
(2009) with 509 kcal per capita per day. Wheat is the second
(216 kcal per capita per day) and beans third (54 kcal per
capita per day). Per capita consumption of beans ranks
second in the region (5.8 kg).
The prevalence of undernutrition is very high at 26%
(201012), a sharp increase from figures of 13% to 17%
recorded in the previous 10 years.
The share of agriculture value added in total GDP was
10.8% in 2012.
The main crop is rice, which accounts for almost all the
harvested grain (paddy rice 220 000 tonnes) according to FAO
statistics for 2012.
Rice availability for consumption (2009 data) was 629 kcal
per capita per day of the milled equivalent. Wheat accounted
for 413 kcal per capita per day and soybean oil for 263 kcal per
capita per day. Cereals share in total food supply was 41.6%.
Uruguay

Located on the southeast coast of South America, most of

Uruguays landscape is a mixture of flat plains and ranges
Peru
of low hills, known as Cuchillas. It has the largest proportion
of arable land (10.3%) and agricultural land (20.8%) in the
The birthplace of the Incas empire is located on the western
region, employing 10.99% of the total labor force in
coast of the continent, facing the Pacific. Peru has three dis-
agriculture.
tinctive environments: the coastal region, the Andes, and the
Uruguay, although small in area, manages to export most of
eastern jungle of the Amazon basin.
its grain and oilseed crop productions. The agricultural sector
Almost one-fourth of the population (23% in 2012) is
as a whole has mimic in many ways the evolution that hap-
dedicated to agricultural activities. There has been a steady
pened in Argentina and the south of Brazil, where contractors
decline in the evolution of rural population at a rate of 1.5%
(mainly from neighboring Argentina) use traditional livestock
per year in the last decade.
lands to produce cash crops such as soybean (1 180 000 hect-
Arable land has remained at around 3 650 000 hectares in
ares sown in 2012) in short-term leases using the latest tech-
the last 10 years, which represents 2.8% of the total area of the
nology (no-till machinery, GMOs, and fertilizers). As a result of
country. Permanent crops have increased in area from
this, soybean production evolved from < 70 000 tonnes in the
500 000 hectares in 1996 to 850 000 hectares in 2011.
year 2000 to 3 000 000 tonnes in 2012.
The share of agricultural value added in total GDP is low,
Wheat and rice are the cereal largest crops, with harvests of
when compared with other countries in the region. In the last
1 000 000 tonnes and 1 490 000 tonnes (paddy), respectively,
decade, agriculture GDP never exceeded 5%.
in 2012. Uruguay is the second producer of barley
The main grain crops are rice (paddy) with a production of
(235 000 tonnes) in the region and the third of sorghum
3 019 316 tonnes in 2012 and maize (1 679 333 tonnes in
(105 000 tonnes).
2012). Quinoas production was 44 000 tonnes (2012).
Exports of soybean (1 810 000 tonnes in 2011) and wheat
Peru produces around 90 000 tonnes year1 (201012) of
(900 000 tonnes in the same year) are the most valuable for
dry beans and exports an average of 34 000 tonnes year1.
the country, only surpassed by boneless meat exports of veal
Peru is a large importer of grains (third in South America).
and beef.
In 2011, maize and wheat imports accounted for
Wheat is the top grain commodity available for consump-
1 907 853 tonnes (the second largest maize importer in the
tion (2009) with 891 kcal per capita per day. Rice (milled) at
region and fifteenth in the world) and 1 683 702 tonnes,
192 kcal per capita per day and maize (177 kcal per capita per
respectively.
day) rice are also important. Cereals account for 45.6% of the
Rice is the top grain commodity available for consumption
total food supply (2009).
(2009) with 524 kcal per capita per day. Wheat is the second
Together with Argentina, Chile, and Venezuela, Uruguay
(345 kcal per capita per day) and maize fifth (184 kcal per
has a consistently low number of the population undernour-
capita per day). Cereals account for 42.9% of the total food
ished. Undernourishment is under 5%, while in 199092,
supply (2009). Peru is one of the largest pasta consumers in the
was 7.3%.
region with 250 000 tonnes year1.
The prevalence of undernutrition is 11% (201012), a
sharp decrease from figures of 23% recorded more than a
Venezuela, Bolivarian Republic of
decade ago in 19992001.
Agriculture is concentrated in the Llanos, a vast tropical grass-
land plain, and does not constitute a major component of the
Suriname Venezuelan economy. Even though there have been domestic
production increases in rice, corn and sorghum in recent years,
Most of the agricultural products are cultivated on reclaimed the country heavily relies on grain imports. Share of agriculture
land in the coastal region. value added in total GDP was 3.9% in 2012.
Almost all the country is covered by forest. Arable land Arable land represents 2.9% of the total country area and
average just 60 000 hectares (0.4% of total area), and rural has remained stable in the last decade. Rural population has
population accounted for 29.78% of total population. dropped from 11.58% in 1997 to just over 6% in 2012.
 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America
Maize (mostly white) is the most important cereal in
Venezuela, constituting the main source of energy in the Ven-
ezuelan diet. Refined, precooked maize flour is the basic ingre-
dient for the preparation of arepas, a popular type of corn
cakes. The availability for consumption in 2009 was 429 kcal
per capita per day. Other top ten grain commodities available
for consumption were wheat (371 kcal per capita per day) and
milled rice (308 kcal per capita per day).
The Venezuelan government is committed to increase the pro-
duction of maize, particularly white. For the period 19922012,
the average production of maize had been 1 709 294 tonnes. Pro-
duction figures for 2011 were over 2 118 000 tonnes and
3 000 000 tonnes for 2012 (FAOSTAT, unofficial data).
Rice (paddy) average production (19922012) was
915 724 tonnes.
Wheat is imported, on average, at a rate of 1 600 000 tonnes
annually. Domestic production is almost inexistent, due to
unsuitable climate and soil conditions.
Pasta and bread are low-cost basic staples of the Venezuelan
diet. In terms of pasta consumption, Venezuela ranks second in
the world (12.3 kg per capita in 2011), behind Italy (26 kg per
capita) (Figure 4).
Beans production was 47 000 tonnes in 2012. Consump-
tion averaged 5.3 kg per capita per year in 2009, the third
largest in South America, behind Brazil and Paraguay. Pabel-
lon is a national dish, which consists of rice, beans, beef, and
fried plantains.
Undernourishment is under 5%, while in 199092,
was 13.5%.
14
12.3
12
10
8.4
kg per capita per year
8.2
8
6
4
0
Venezuela Chile Peru
Figure 4 Pasta consumption in South American countries (2011). Source: www.internationalpasta.org.
427
Future Prospects
Brazil and Argentina are likely to maintain and even expand
their leadership in grain production and trade in the region.
Spare farmland will continue to become available for
increasing grain and oilseed crop production, particularly in
El Cerrado, Brazil. According to USDA, Brazil will duplicate its
soybean harvest in 10 years, but they will need to solve urgent
infrastructure shortcomings.
Yield gains are still possible in the region, continuing with
the success achieved in the last decades. This would not be
restricted exclusively to large farms; small and medium pro-
ducers may also benefit from improved efficiency in the man-
agement of their production systems.
No-tillage technology and GMOs together with adequate
cultural practices have proved to be key factors for the success
of the agricultural sector in the region.
All the previously mentioned factors built on the bases
of a very strong agricultural research and technology sector,
both government-driven and private. Nevertheless, more
actions are needed to secure sustainability of the model.
For example, broadening the use of irrigation would help
to increment agricultural productivity and crop
diversification.
On the matters and nutrition and food security, FAO and
Mercosur have joined forces to strengthening family farming in
the region, which accounts for nearly 40% of Brazil agricultural
production,30% in Uruguay, 25% in Chile, 20% in Paraguay,
and 19% in Argentina.
7.9
6.2
4.8
3.9
3
Argentina Brazil Bolivia Ecuador Colombia
428 GRAINS AROUND THE WORLD | Grain Production and Consumption: South America
Climate changes in the region may negatively impact from
an economic and social perspective if not properly addressed.
Exercises for Revision
What pseudocereals are native to South America?
What is the social cost of the agricultural economic boom in
the region?
What strategy is Brazil using to mitigate hunger in its
population?
Colombia is one of the major importers of maize in the
region. What is it mainly used for?
How is Ecuador planning to increase its maize production?
Why are breakfast cereals not popular in the region?
Where is El Cerrado located and what is its importance?
Name the countries where wheat is not the main staple
cereal in the region.
Find a recipe for Feijoada and Pabellon on the Net. What
grain components do they share?
What were the main economic and political reasons behind
the 2008 farmers strike in Argentina?
Exercises for Readers to Explore the Topic Further
Sustainability of the soybean production models
Brazil agriculture before and after lifting the ban on GMOs
No-tillage and postharvest technology in the region
See also: Agronomy of Grain Growing: Precision Agriculture;
Soybean: Agronomy; Wheat: Agronomy; Non-wheat Foods: Maize:
Foods from Maize; The Legumes and Pseudocereals: Amaranth:
Overview; Quinoa: Overview; The Oilseeds: Oilseeds: Overview;
Wheat-Based Foods: Breads; The Cereal Grains: An Overview of
the Family of Cereal Grains Prominent in World Agriculture.
Further Reading
Cereal marketers race for global bowl domination http://adage.com/article/news/cereal-
marketers-race-global-bowl-domination/236738/ (accessed 17/12/13).
Quinoa boom a mixed blessing for Bolivians http://www.aljazeera.com/indepth/features/
2013/07/201371013540775237.html (accessed 12/12/13).
Brazil says yes to genetically modified food, Mexico says no http://www.
businessweek.com/articles/2013-10-30/brazil-says-yes-to-genetically-modified-
foods-dot-mexico-says-no (accessed 12/12/13).
Nutrition Profile Bolivia http://www.childinfo.org/files/nutrition/DI%20Profile%20-%
20Bolivia.pdf (accessed 12/12/13).
Nutrition Profile Paraguay http://www.childinfo.org/files/nutrition/DI%20Profile%20-%
20Paraguay.pdf (accessed 15/12/13).
Why is Argentina so efficient? http://cornandsoybeandigest.com/tillage/why-argentina-
so-efficient (accessed 15/11/13).
Dros JA (2004) Managing the soy boom: Two scenarios of soy production expansion in
South America. Amsterdam: AIDEnvironment.
The miracle of the cerrado http://www.economist.com/node/16886442
(accessed 17/12/13).
Ecuador 2013 maize production estimated at record levels http://www.thecropsite.com/
news/15111/ecuadors-2013-maize-production-estimated-at-record-levels
(accessed 4/1/14).
Current status of adoption of no-till farming in the world and some of its main benefits
http://www.fao.org/ag/ca/CA-publications/China_IJABE.pdf (accessed 14/11/13).
Brazil tries to fill the potholes in its path to growth http://www.ft.com/cms/s/0/
7dd98ed6-059f-11e3-ad01-00144feab7de.html#axzz2xV094L00
(accessed 31/3/14).
Pasta Consumption http://www.internationalpasta.org (accessed 1/11/13).
Family Agriculture in the Mercosur http://www.reafmercosul.org/reaf
(accessed 13/12/13).
Sunny side up for Latin America breakfast cereals http://www.just-food.com/analysis/
sunny-side-up-for-latin-america-breakfast-cereals_id97594.aspx (accessed 17/12/
13).
GMOs a global debate: Brazil, second largest GMO producer in world http://www.
theepochtimes.com/n3/162906-gmos-a-global-debate-brazil-second-largest-gmo-
producer-in-world/?photo2 (accessed 18/12/13).
Value Added (%GDP) in Colombia http://www.tradingeconomics.com/colombia/
agriculture-value-added-percent-of-gdp-wb-data.html (accessed 12/12/13).
Vergara W, Rios A, Trapido P, and Malarin H (2014) Agriculture and future climate in
Latin America and the Caribbean: Systemic impacts and potential responses. Inter-
American Development Bank Discussion Paper No. IDB-DP-329.
Relevant Websites
http://www.faim.org.ar/ Argentinean Milling Industry Federation (In Spanish).
http://odepa.gob.cl Chilean Office of Studies and Agricultural Policies (In Spanish).
http://www.embrapa.br/english Embrapa (Brazilian Agricultural Research
Corporation).
http://www.reafmercosul.org/reaf/ Family Agriculture in the Mercosur.
http://faostat.fao.org/site/291/default.aspx FAOSTAT.
http://inta.gob.ar/ INTA (Argentinean National Agricultural Technology Institute) (in
Spanish).
http://www.iica.int/Eng/Pages/default.aspx Inter-American Institute for Cooperation
on Agriculture.
http://www.thecropsite.com/reports/?id2556 The Crop Site.
http://www.rlc.fao.org/en/publications/zero-hunger/ The Fome Zero (Zero Hunger)
Programme.
 Scientific Organizations Related to Grains
Contents
Research Organizations of the World: CGIAR
Research Organizations of the World: Europe and North America
Research Organizations of the World: Asia-Pacific, CentralSouth America, and AfricaMiddle East
Research Organizations of the World: Global Trends and the Commercial Sector
Scientific Societies Associated with Grain Science
Research Organizations of the World: CGIAR
JH Skerritt, University of Queensland, St Lucia, Australia
2016 Elsevier Ltd. All rights reserved.
This article includes some content from the previous edition article by F.J.B Reifschneider & S. Hussain, volume 3, pp 2636, 2004, Elsevier Ltd.
Topic Highlights
The CGIAR international agricultural research centers pro-
vide a significant contribution to grains research, particu-
larly for the developing world.
The CGIAR centers have traditional strengths in germplasm
development, agronomy, quality assessment, biotechnol-
ogy, pathology, natural resource management, and
economics.
In 201112, much of the research undertaken within the
CGIAR system was organized into major research programs.
Five CGIAR Research Programs have a primary emphasis on
improvement of yields and profitability of grain crops,
dryland cereals, grain legumes, maize, rice, and wheat,
while other programs focus on farming systems, crop nutri-
tional characteristics, production sustainability, soil and
water management, and climate change.
Learning Objective
To identify and understand the role and major priorities of
the CGIAR, particularly its centers that focus on grains
research.
Current and Future Challenges
Agriculture is inextricably linked to the broader development
agenda. Most of the worlds poor people depend directly or
indirectly on agriculture for their livelihoods although almost
one billion people in the world are undernourished. The
Earths ecosystems are under stress: Almost half of land has
already been irreversibly transformed or degraded by human
actions and agricultural activities. Most of the worlds water is
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00055-3
used for irrigation, and agriculture frequently accounts for 70%
of freshwater withdrawals and agricultural activities are one of
the leading causes of climate change, contributing 20% of all
greenhouse gases.
Globally, the agricultural sector is set to face pressing chal-
lenges in the future:
World population is expected to reach nine billion by 2050.
Over the same time frame, demand for food is set to more
than double, with most of the increased demand coming
from developing countries.
Erosion, salinization, compaction, and other forms of soil
degradation affect 30% of the worlds irrigated lands, 40%
of rainfed agricultural lands, and 70% of rangelands.
Most of the worlds freshwater is used for irrigation and
water use is expected to increase by 50% by the 2030s.
Arable land per person in developing countries was 0.32 ha
in 196163, but is expected to drop to 0.16 ha by 2030.
Cereal productivity growth increases (i.e., rises in cereal
yields per ha) have slowed.
Given these challenges, science-based agricultural develop-
ment will continue to be an important method for promoting
economic growth and addressing current and future food and
environmental challenges and poverty. Concern about hunger
and the worlds ability to feed growing populations has been a
recurring theme throughout history. In modern times, the Irish
famine and the famines of the twentieth century in Bengal
(194344), Bangladesh (197475), China (195961), and
Holland (194445) and during the 1970s80s in large parts
of Africa (Ethiopia, Sudan, Mozambique, Nigeria, Niger,
Somalia, and Zaire) caused millions of deaths. Famines can
result from breakdowns in food grain production and
distribution and from a restricted flow of income with which
people buy food. Shocks in the agricultural sector (plant dis-
ease as in the case of Irish famine and extreme weather events
429
430 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR
such as droughts and floods) and human-induced conflicts
and civil wars can all trigger famines.
The 1960s marked a period when deep-rooted pessimism
about food availability surfaced anew. However, scientists
working with farmers successfully averted the worst Malthu-
sian specter by ensuring abundant agricultural yield. The story
of wheat improvement in England is compelling. It took nearly
1000 years for wheat yields to increase from 0.5 to 2 tonnes
per hectare (mt ha 1), but only 40 years to climb from 2 to
6 tonnes ha 1, illustrating the ability of modern grain science
to increase agricultural productivity.
Mobilizing Grain Science for Development
The Consultative Group on International Agricultural Research
(CGIAR) was established in 1971. A group led by The Rocke-
feller Foundation, Ford Foundation, Food and Agriculture
Organization, The World Bank, and others created the
CGIAR. The founding objective was based on a profound belief
in the ability of science, especially grain science, to increase
food supply in low-income countries facing severe food scar-
city. At that time, the central idea of mobilizing grain science
internationally to meet food and development needs, within a
public goods framework, was novel.
The CGIAR system is supported by the World Bank and a
range of national governments, usually through their overseas
development assistance programs. In 2013, the CGIARs
research investment exceeded $US 1 billion for the first time
and represented the single largest effort in mobilizing science
for agricultural development. The CGIAR is best known for the
green revolution, a term used to refer to the set of agricultural
innovations (new high-yielding varieties of wheat, rice, and
maize, increased investments in irrigation, easy availability of
farm inputs such as fertilizers and pesticides, improved grain
distribution, and food policies in developing countries) that
helped boost grain yields and incomes of farmers in poor,
agrarian economies. The green revolution transformed agricul-
ture in developing countries especially in India, Mexico,
Pakistan, and the Philippines.
Cereal crop yields and food production more than doubled
between 1960 and 1985. The new varieties of rice, wheat, and
maize increased daily calorie supply in developing countries by
25%, from under 2000 calories per person in the early 1960s to
2500 calories per person by the 1990s. Without the productiv-
ity growth in basic food crops, the number of poor and hungry
would have been far greater.
Critics of the green revolution have argued that the new
crop technologies favored farmers in well-endowed areas,
worsened pollution through excessive use of fertilizers and
pesticides, threatened biodiversity, promoted monocultures,
and required higher input costs from impoverished farmers.
While a comprehensive discussion of these issues is beyond the
scope of this article, subsequent studies and analyses have
increased understanding about the effects of the new technol-
ogies. A range of subsequent policy interventions (e.g.,
increased credit, improved marketing and seed distribution
facilities, land reforms, and strengthened emphasis on pesti-
cide safety, and others) have helped poor farmers to benefit
from green revolution technologies.
However, the benefits of the green revolution were spread
unevenly and largely bypassed Africa. On average, crop yields
in Africa are one-third of those achieved by Asian farmers. Less
than 4% of Africas arable land is irrigated. There is enormous
potential for growth-led productivity in sub-Saharan Africa
where agriculture provides 70% of employment, 40% of
exports, and one-third of gross domestic product. However,
productivity benefits for Africa are now starting to emerge,
particularly for maize, roots, and tubers.
Grains Research Centers in the CGIAR
CGIAR achieves its mission through the work of 15
international agricultural research centers (IARCs, Table 1)
that work with national agricultural research systems, the pri-
vate sector, and civil society. The IARCs are autonomous insti-
tutions, linked by a common commitment to promoting
agricultural development in developing countries; 13 IARCs
are headquartered in developing countries.
Improving the sustainable productivity of food grains
(including cereals, legumes, and pulses) is a major objective
of CGIAR research. Nine of the IARCs have specific crop
improvement mandates. More specifically, the CGIAR centers
have traditional strengths in germplasm development, provid-
ing much of the germplasm for development of the modern
varieties that are now utilized throughout most of the devel-
oping world (as well as in some developed countries with
similar production environments such as Australia). To sup-
port this work, centers have strengths in agronomy, quality
assessment, biotechnology, pathology, natural resource man-
agement, and economics.
For the purposes of this paper, only eight IARCs have crop
improvement mandates that correspond to the definition of
grains cited earlier, the exception being root and tuber crops,
including potato. Even though Bioversity (formerly known as
International Plant Genetic Resources Institute) does not have
a plant breeding program, its mandate to serve as a world
center for conserving plant genetic resources is germane to
the focus of this article.
The International Agricultural Research Centers
Several of the centers have a grain science focus, including the
Centro Internacional de Mejoramiento de Maz y Trigo (The
International Maize and Wheat Improvement Center
(CIMMYT) www.cimmyt.org). CIMMYT is based in Mexico
but has programs in Latin America, Africa, Central and West
Asia, and North Africa. Three centers carry out studies on rice
improvement. The International Rice Research Institute (IRRI,
www.irri.org) is the main one, and its work aims to develop
new rice varieties and crop management techniques to help
farmers improve the yield and quality of rice in a sustainable
way. The semidwarf rice varieties and management packages
developed by IRRI were central to the successes of the green
revolution in the 1960s and 1970s. Two other centers under-
take rice breeding and management in Latin America and
Caribbean (International Center for Tropical Agriculture
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR
Table 1 CGIAR Centers with crop improvement mandates
Year
Center founded
International Center for Tropical Agriculture (CIAT) 1967
International Maize and Wheat Improvement Center 1966
(CIMMYT)
International Center for Agricultural Research in the Dry 1975
Areas (ICARDA)
International Crops Research Institute for the Semi-Arid 1972
Tropics (ICRISAT)
International Institute of Tropical Agriculture (IITA) 1967
International Plant Genetic Resources Institute (now called 1974
Bioversity)
International Rice Research Institute (IRRI) 1960
International Potato Center (CIP) 1970
West Africa Rice Development Association (now called 1975
AfricaRice)
a
Currently Lebanon.
b
Currently Benin.
(CIAT) ciat.cgiar.org in Cali, Colombia) and West Africa
(AfricaRice, www.africarice.org, in Cotonou, Benin).
Other CGIAR centers with a strong grains focus include the
International Crops Research Institute for the Semi-Arid Tro-
pics (ICRISAT, www.icrisat.org) in Patancheru, India (focussing
on sorghum, pearl millet, chickpea, pigeon pea, and peanut),
and the International Center for Agricultural Research in the
Dry Areas (ICARDA, www.icarda.cgiar.org) long established in
Syria, but now headquartered in Beirut, Lebanon (focussing on
barley, wheat, lentil, faba bean, chickpea, and forage legumes).
The CGIAR centers have traditional strengths in germ-
plasm development, providing much of the germplasm for
development of the modern varieties that are now utilized
throughout most of the developing world (as well as in some
developed countries with similar production environments
such as Australia). To support this work, centers have
strengths in agronomy, quality assessment, biotechnology,
pathology, natural resource management, and economics. At
CIMMYT, wheat breeding and maize breeding (agronomy
and quality evaluation programs) are backed up by programs
in economics, natural resource management, and biotechnol-
ogy. ICRISAT has a geographic emphasis on South Asia and
sub-Saharan Africa, and its research addresses water scarcity
through natural resource management (rainwater conserva-
tion and utilization) and plant breeding to improve water-use
efficiency and drought tolerance. ICARDA has a geographic
emphasis on central West Asia and North Africa, but its germ-
plasm is utilized worldwide. ICARDA focuses on agriculture
in tropical dry areas, while ICRISAT emphasizes temperate
and subtropical dry areas. It also has an emphasis on water-
soil management in integrated farming systems for low-
rainfall areas. The International Institute of Tropical Agricul-
ture (IITA) with headquarters in Nigeria focuses on germ-
plasm (including cowpea, maize, and soybean), pest
management, and crop-based farming systems for small-
holder farmers of sub-Saharan Africa. Some work on proces-
sing quality is carried out by the centers, but little work on
grain storage and other postharvest technologies.
431
Host
country Mandate crops
Colombia Cassava, beans, rice, tropical forages
Mexico Maize, wheat (bread and durum wheat, triticale)
Syriaa Barley, wheat (durum species), lentil, chickpea, faba bean,
forage legumes
India Sorghum, pearl millet, groundnut, chickpea, pigeon pea
Nigeria Cassava, yams, maize, soybean, cowpea
Italy Genetic resources, cocoa, coconut/Musa species
Philippines Rice
Peru Potato, sweet potato
Cote Rice
dIvoireb
Evolving Research Agenda
The CGIARs role and mission has evolved over time and this is
reflected in its structure. When the CGIAR was founded, four
IARCs (CIAT, CIMMYT, IITA, and IRRI) were brought under its
umbrella. Thus, at that time, it was largely an alliance of grains
research organizations, and in the 1960s, the research agenda
was clearly around maize and rice productivity.
During the 1970s, new IARCs were established to pioneer
improvements in key food crops (legumes, roots, tubers, and
other cereals), livestock health, and ecological regions (dry, semi-
arid, and tropical regions). In the 1980s, the mandate (and struc-
ture) was expanded to encompass food policy and strengthening
of national agricultural research systems and in the 1990s further
expanded to conduct research on pressing issues involving agro-
forestry, forestry, water management, and aquatic resources.
During the 1990s and early-mid-2000s, the research pro-
grams undertaken in the CGIAR were organized around five
focus areas:
Increasing productivity (of crops, livestock, fisheries, for-
estry, and the natural resource base)
Strengthening national systems (joint research, policy sup-
port, training, and knowledge sharing)
Protecting the environment (by developing new technologies
that make more prudent use of land, water, and nutrients and
help reduce agricultures adverse impacts on ecosystems)
Saving biodiversity (collecting, characterizing, and conserv-
ing genetic resources)
Improving policies (with a major impact on agriculture,
food, health, the spread of new technologies, and the man-
agement and conservation of natural resources).
In summary, over 50 years, the CGIARs research portfolio
evolved from the original focus on increasing productivity in
individual critical food crops. The current approach recognizes
that saving biodiversity, protecting the environment, improving
food policies, and strengthening scientific capacities of develop-
ing countries through training and knowledge sharing are all key
432 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR
components in the drive to enhance sustainable agricultural
productivity. The CGIARs focus on the fundamentals remains
as strong as ever: Agricultural growth and increased farm pro-
ductivity in developing countries creates wealth, reduces poverty
and hunger, and protects the environment.
Recognizing the complexity of the challenges faced in devel-
oping world agriculture and the importance of establishing
sufficient capacity, the CGIAR was reorganized effective 2010.
The CGIAR Consortium (www.cgiar.org/cgiar-consortium)
integrates the research of the 15 centers to increase the effec-
tiveness and relevance of the work of the CGIAR; enhance the
impact of the research by setting common objectives and plan-
ning concerted action; and to take advantage of opportunities
for the CGIAR to become more relevant, more effective, and
more efficient.
In 201112, much of the research undertaken within the
CGIAR system was organized into major research programs
(www.cgiar.org/our-research/cgiar-research-programs),
which typically involve several CGIAR centers as well as uni-
versities, NGOs, and government agencies from both devel-
oping and developed countries. The CGIAR Consortium
coordinates CGIAR Research Programs across research
centers, which are members of the CGIAR Consortium and
partner.
Five CGIAR Research Programs have a primary emphasis on
improvement of yields and profitability of grain crops:
Dryland cereals (such as barley, sorghum, finger, and pearl
millets) in South Asia and Africa
Grain legumes (addressing major biotic and abiotic stres-
ses, nitrogen fixation, and opportunities to use grain
legumes to intensify cropping systems)
Maize (focussing on the productivity of maize-based crop-
ping systems)
Rice (focussing on genetic diversity, development of
new varieties, rice-based farming systems, quality, and
marketing)
Wheat (wheat-based cropping systems, pest, diseases, and
nutrient management, yield, and seed systems)
In addition, two programs (Integrated Systems for the Humid
Tropics and Dryland Systems) aim to improve the productivity,
profitability, sustainability, and resilience of entire farming
systems; one program (Agriculture for Nutrition and Health)
includes a thrust on grains nutritional composition, while a
further program underpins the CGIAR crop (particularly grain)
germplasm collections. Further two programs aim to improve
production sustainability and environmental integrity and
adapt to and mitigate climate change namely, the CGIAR
Research Programs on Climate Change, Agriculture and Food
Security and Water, Land and Ecosystems.
Collaboration with National Agricultural Research
and Extension Systems
As comparatively small, research-oriented centers focussed on
international public goods, the tension between addressing
this mission and achieving practical impacts at the national
level and in the shorter term has been present since the
formation of the CGIAR and its forerunners over 50 years
ago. Another continuing challenge is the level of skills and
resourcing within the National Agricultural Research and
Extension Services (NARES) as this affects the ability of
national organizations in poorest countries to be able to effec-
tively partner. Centers with an ecoregional focus, such as
ICARDA, have traditionally had a more extensive program of
collaboration with the NARES in their region than those with a
global crop mandate. The NARES themselves have recognized
the importance of leveraging off CGIAR research and trying to
do so in a more structured manner than in the past. For
example, the Asia-Pacific Association of Agricultural Research
Institutions (APAARI) has recently developed a strategy for
strengthening partnerships in South Asia between the NARES
and the CGIAR (www.apaari.org/publications/nars-cgiar-pro-
ceedings.html).
Various approaches have been used to develop or
strengthen relationships between centers and the NARES in
developing countries. One of the most effective and enduring
has been development of mechanisms by each of the grains
centers to ensure that developing country breeders are trained
at their centers and that grains breeding programs have access
to materials. This has typically involved provision of germ-
plasm and advanced lines (but not usually finished varieties),
although the level of crossing from the materials made avail-
able by the centers depends on the resources available in the
particular NARES as well as the extent of further work required
to ensure that the material is sufficiently locally adapted.
Translation (or extension) of agronomic research results
from CGIAR centers to the NARES is necessarily more
complex by definition, it involves practice change rather
than having the technology embedded in the seed. Apart
from the expected approaches of short- and long-term training
and development of publications and other written materials,
a challenge has been to develop longer-term partnerships that
achieve impact while not distracting from the core function of
knowledge generation at the centers.
To facilitate such partnerships, a number of individual
national and multilateral donors have funded major partner-
ship projects, which through the provision of restricted
funding (now called bilateral funding) can fund the exten-
sion of CGIAR crops research results through formal, funded
partnership projects. For example, the African Development
Bank is currently supporting a major collaboration between
AfricaRice, ICARDA, and IITA to enhance the productivity and
income of four priority value chains (cassava, maize, rice, and
wheat) on a sustainable basis. The importance of partnerships
with the NARES has also been recognized in the design of
most of the new CGIAR Research Programs and most projects
that involve not only several CGIAR centers but also the
NARES and universities in both developed and developing
countries.
Finally, capacity building of the NARES organizations has
itself been a focus of the CGIAR over many years. This led to
the formation of a specialized CGIAR center, the International
Service for National Agricultural Research (ISNAR, in the
Netherlands), although it merged into the IFPRI in 2004.
Some controversy remains, however, as to whether the
CGIAR is weakening the NARES capacity rather than building
it as in some regions the most able, internationally trained
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR
scientists are hired to work for the CGIAR institutes. In addi-
tion, where the NARES are weak, there has in the past been a
tendency for the CGIAR to assume the NARES role of develop-
ing and testing technologies at plot and field scale. This returns
us to the challenge of demonstrating impact versus seeing the
CGIAR as an intermediary.
Selected Impacts of CGIAR Grains Research
There have been several published analyses of the overall
impacts of CGIAR research, and most of them have shown an
overwhelmingly positive return on investment. In the initial
studies, the point was made that in some countries, the bene-
fits from modern varieties developed by the CGIAR centers had
favored poor urban food consumers and large farmers rather
than poor smallholder farmers, although it was pointed out
that small farmers do tend to adopt modern varieties but later
than large farmers. Evenson and Gollin published an extensive
study of CGIAR crop research impacts across a wide range of
crops and continents. Renkow and Byerlee assessed the
impacts of more recent GIAR research that published since
2000 and argued that impacts of CGIAR crops genetic
improvement research have continued to be strongly positive
even some decades after the green revolution. Lastly, Raitzer
and Kelley attempted to undertake a benefitcost meta-analysis
of investment in CGIAR research and concluded benefitcost
ratios of between 1.9 and 17.3, depending on the stringency of
demonstration of causality of benefits from particular research
activities. However, they concluded that the true benefits
arising from the CGIAR is probably in excess of even the
upper bounds of these results as only a small subset of system
impacts have been quantified.
Wheat
CIMMYT has a global mandate for wheat improvement (bread
and durum species). In the late 1960s, about one-third of all
wheat varieties in developing countries were CIMMYT crosses.
By the 1990s, these numbers rose to about half from CIMMYT
crosses and another quarter from varieties that had a CIMMYT
parent. Worldwide, around 90% of all spring bread wheat
releases had at least one CIMMYT ancestor; the percentage for
durum wheat was even higher with nearly all spring durum
wheat releases having a CIMMYT ancestor. In 1970, semidwarf
wheat varieties were important only in Asia. By the 1990s,
semidwarf wheat varieties covered 80% of world wheat area,
with adoption rates of 90% and higher in Asia and Latin
America. More recently, across Afghanistan, Bangladesh,
Ethiopia, Egypt, Nepal, and Pakistan, high-yielding wheat vari-
eties resistant to Ug99, a highly virulent disease, have protected
the livelihoods and food security of 500 000 farming families.
ICARDAs wheat improvement research focuses on spring
bread and durum wheats in the West AsiaNorth Africa region.
ICARDA grain scientists have made major advances in identi-
fying new sources of resistance to abiotic stresses (drought,
heat, and cold) and biotic stresses (rusts, Septoria diseases,
and Hessian fly), in broadening genetic diversity and in select-
ing for grain quality. ICARDAs genebank holds over 135 000
accessions from over 110 countries: traditional landraces,
433
improved germplasm, and a unique set of wild crop relatives.
These include wheat, barley, oats, and other cereals; food
legumes such as faba bean, chickpea, lentil, and field pea;
and forage crops, rangeland plants, and wild relatives of each
of these species.
Rice
IRRI began rice improvement in the 1960s. Expanding on the
successes in wheat improvement, IRRI scientists successfully
introduced semidwarfism into two indica rice varieties (IR5
and IR8). The latter variety would lay the foundations of the
green revolution in Asia. IRRI has made substantial contribu-
tions toward improvement of rice varieties. Of the estimated
2040 rice varieties released over 40 years by national rice
research systems of South and Southeast Asia, 219 were direct
IRRI lines. Of the 2021 varieties, 31% had originated from one
or more parents developed at IRRI. This number rises to 46%
when the IRRI-developed varieties are included. In the 1970s,
the release of IRRI-related varieties peaked at 60%, subse-
quently stabilizing at 40% of all varieties released. Since that
time, the increase in rice production has been enormous. More
recently, Scuba rice, which can survive under water for 2 weeks,
is protecting the harvests, incomes, and food security of poor
farmers and consumers across monsoon Asia.
AfricaRice conducts rice improvement research in West and
Central Africa, focussing on three distinct rice ecologies:
upland, rainfed lowland, and irrigated land. Using interspecific
hybridization techniques, Africa Rice successfully combined
the ruggedness of the local African species of rice (Oryza gla-
berrima) with the phenomenally high productivity traits of
Asian rice (O. sativa) (common rice) that was the mainstay of
the green revolution. The new rice varieties, NERICA or New
Rice for Africa, resist droughts and pests and are able to thrive
in poor soils. CIAT has a mandate to promote sustainable
increases in rice production and productivity in Latin America
and the Caribbean region.
Maize
CIMMYT has a global mandate for maize improvement, and it
targets lowland tropical, subtropical, midaltitude, and tropical
highland environments throughout the developing world.
Mexico, where CIMMYT headquarters are located, is a center
of origin for maize. The use of CIMMYT germplasm has
increased over time, with about two-thirds of public sector
varietal releases containing CIMMYT germplasm. Estimates of
economic benefits from CIMMYTs maize research vary, but
based on the earlier results (21.2 Mha planted to CIMMYT-
related varieties) and a conservative estimate of farm-level
maize prices (US$120 per tonne, on import parity basis),
gross benefits range from US$1.3 billion to US$4 billion per
year. These benefits take into account both germplasm
improvements and better crop management practices.
CIMMYT also developed quality protein maize (QPM), a
product of 30 years of research. QPM looks and tastes like
normal maize but has nearly twice the amount of lysine and
tryptophan, two amino acids essential for protein synthesis in
humans. QPM is now being planted on 1 Mha in 20 countries,
boosting food, nutrition, and income security. More recently,
434 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR
drought-tolerant maize has increased farmers yields by
2030%, benefiting 20 million people in 13 African countries.
IITA has a regional maize improvement mandate mainly tar-
geting humid tropical and moist savannah zones of Western
and Central Africa. Maize accounts for more than 20% of
domestic food production in Africa.
Other Crops
Two significant achievements involving ICRISAT should be
mentioned. Breeding and disseminating micronutrient-rich
crop varieties through biofortification are potentially a cost-
effective strategy to increase micronutrient intake such as iron
and zinc. India has some of the worlds worst malnutrition rates,
resulting from a lack of essential vitamins and minerals in the
diet. This affects the survival and development of women and
children in particular. About three quarters of children under
three and half of women suffer from anemia, mostly caused by
iron deficiency. Pearl millet is a nutritious but underutilized
staple crop, eaten by 50 million people in dry rural areas of
India. As well as being rich in B vitamins, calcium, iron, potas-
sium, magnesium, and zinc, it is well adapted to drought, poor
soils, and high temperatures. It is much higher in iron than
wheat, rice, or maize. ICRISAT is developing drought-tolerant
pearl millet varieties with even higher levels of iron. This is part
of a larger biofortification initiative coordinated through Har-
vestPlus and IFPRI and reviewed in Saltzman et al.
Another significant impact involves partnership with agri-
business on the microdosing of fertilizer to resource-poor
farmers in sub-Saharan Africa. This involves using only
810 kg of nitrogen per hectare but applying it precisely to the
roots three to 4 weeks after planting. In Zimbabwe, ICRISAT has
recorded yield increases of 3050% among farmers adopting the
technique. With evidence that precision agriculture techniques
can work, the challenge is creating appropriate enabling envi-
ronments to encourage take-up. In addition, ICRISAT has been
active in developing agribusiness incubators, but the involve-
ment of other centers in agribusiness has been variable.
Conserving Biodiversity
For more than 30 years, CGIAR scientists have been collecting,
characterizing, and conserving biodiversity. Currently, the
CGIAR holds over half a million samples of crop, forage, and
agroforestry genetic resources in 11 gene banks around the
world. Since the 1980s, CGIAR gene banks have distributed
well over one million germplasm samples worldwide, and the
vast majority of requests came from developing countries. In
1994, CGIAR signed an agreement with FAO, placing the col-
lections in public trust, available to researchers worldwide.
Recognizing the challenge of creating a sustainable funding
base for conservation activities, the Global Crop Diversity Trust
(GCDT) was established in association with the CGIAR to
support global conservation of biodiversity. More recently,
the Svalbard Global Seed Vault has been established as a secure
seedbank in Norway, only 1300 km from the North Pole. It
maintains duplicate samples of seeds held in other genebanks
worldwide, as an attempt to insure against the loss of seeds in
other genebanks during large-scale crises.
The CGIAR, with greater emphasis on the centers working
together with each other and with partners in developed and
developing countries to leverage complementary skills and
resources expect a number of significant impacts from the
new CGIAR Research Programs. Some of the planned impacts
from the new programs related to grains (www.cgiar.org/our-
research) include the following:
By 2035, research on rice will increase farmers yields and
lower prices for poor consumers, lifting 150 million people
out of poverty and reducing the number of undernourished
people in Asia by 62 million.
By 2020, 12 million households in Africa will have access to
sustainable irrigation, thanks to research on water, land,
and ecosystems.
By 2022, research will help increase harvests of grain
legumes a key source of protein for the poor in low-
income countries in five regions, improving their nutrition
from 2.1 million tonnes of extra protein.
By 2018, 50 million people will have access to staple food
crops specifically bred to be rich in key vitamins and
minerals namely, iron, zinc, and vitamin A in an effort
to combat malnutrition.
Exercises for Revision
Which CGIAR centers have a mandate for rice research?
What has been the main shift in research organization in
the CGIAR centers over the last 35 years?
What are some of the approaches that have been used to
facilitate the extension of research results from CGIAR cen-
ters to the NARES?
What has been implemented to provide a back up in the
event of a CGIAR centers genebank being destroyed
through war or civil unrest?
Exercises for Readers to Explore the Topic Further
What are the main research activities of IRRI? How are they
funded and what is their relationship with major research
organizations in the developed world?
How do developed countries benefit from the results of
CIMMYT research?
Which centers provide the enabling science in natural
resources management to underpin the work of the crops-
specific centers? How do they collaborate to support each
other?
Acknowledgments
Some of the text, particularly in the Current and Future
Challenges and Mobilizing Grain Science for Development
sections, has been adapted from an article for the first edition
of the encyclopedia, by F.J.B. Reifschneider and S. Hussain,
formerly of the CGIAR office in Washington DC.
See also: Agronomy of Grain Growing: Implication of Climate
Changes; Wheat: Agronomy; Breeding of Grains: Barley: Genetics
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: CGIAR
and Breeding; Maize: Breeding; Rice: Breeding; Soybean: Germplasm,
Breeding, and Genetics; Wheat Breeding: Exploiting and Fixing Genetic
Variation by Selection and Evaluation; Food Grains and the
Consumer: Cultural Differences in Processing and Consumption;
Grains Around the World: Grain Production and Consumption:
China and South-East Asia; Grain Production and Consumption in
Oceania: Australia and Pacific Countries; Grain Production and
Consumption: Overview; Grain Production and Consumption: South
America; Grain Production and Consumption: Africa; Production and
Consumption of Grains: India; Scientific Organizations Related to
Grains: Research Organizations of the World: Asia-Pacific,
CentralSouth America, and AfricaMiddle East; The Basics: The
Grain Crops: An Overview; The Cereal Grains: An Overview of the
Family of Cereal Grains Prominent in World Agriculture; Wheat-
Based Foods: Cereals: Overview of Uses: Accent on Wheat Grain.
Further Reading
Alston JM and Pardey PG (1996) In: Making Science Pay The Economics of
Agricultural R&D Policy, p. 203. Washington, DC: American Enterprise Institute.
Baum WC (1986) Partners Against Hunger: Consultative Group on International
Agricultural Research. Washington, DC: The World Bank.
Binswanger H and McCalla A (2010) In: Pingali P and Enenson R (eds.) Handbook of
Agricultural Economics, pp. 35713712. Amsterdam: Elsevier.
Bouman BAM, Hengsdijk H, Hardy B, Bindraban PS, Tuong TP, and Ladha JK (2002)
Water-wise rice production. In: Proceedings of the International Workshop on
Water-wise Rice Production, 811 April. Los Banos, Philippines: International Rice
Research Institute.
Brown LR (1995) Who Will Feed China? Wake-up Call for a Small Planet. Washington,
DC: Worldwatch Institute.
Bruinsma J (2009) The resource outlook by 2050: How much do land, water and crop
yields need to increase by 2050? Expert Meeting on How to Feed the World in 2050,
Food and Agriculture Organization of the United Nations, ftp://ftp.fao.org/docrep/
fao/012/ak971e/ak971e00.pdf.
435
Conway G (1999) The Doubly Green Revolution Food for All in the 21st Century.
London: Penguin Books.
Evenson RE (2003) In: Evenson RE and Gollin D (eds.) Crop Variety Improvement and
its Effect on productivity The Impact of International Agricultural Research,
pp. 71108. Wallingford, UK: CABI Publishing.
Hazell P (2003) Green revolution. In: Mokyr J (ed.) The Oxford Encyclopedia of
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UK: Cambridge University Press.
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Research and the Poor. CGIAR Study Paper Number 2, Washington DC, USA: The
World Bank.
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109: 1230212308.
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 Research Organizations of the World: Europe and North America
JH Skerritt, University of Queensland, St Lucia, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
While maintaining a systems and sustainability focus, there
has been renewed emphasis on research directed at improv-
ing grain yields in many countries.
The formation of networks between grain research investors,
research institutes, and industry increasing in Europe and
North America.
A greater proportion of agricultural (including grains)
research is carried out in universities in Europe and North
America compared with developing countries, where it is
largely within government-funded research institutes.
Significant increase in commercial involvement in grain
breeding and biotechnology, particularly where strong intel-
lectual property protection and end point royalty systems
are in place. Farmer organization-led grain research is also
becoming more important.
Research to underpin grain utilization for animal feed and
for biofuels is important in several countries.
Learning Objectives
To understand the structure and major roles of public sector
grain research organizations within Europe and North
America.
Introduction
Compared to the long history of cultivation of grains (many
thousands of years), the use of research principles to address
constraints is a comparatively recent approach. Agricultural
societies were established in the United Kingdom, France,
and Germany in the mid-1800s (e.g., The Royal Agricultural
Society of England (1839), La Societe des Agriculteurs de
France (1850), and Deutsche Landwirtschafts-Gesellschaft
(1885)) and by the mid-to late nineteenth century, agriculture
experiment stations were appearing in Europe and North
America. The tradition of government-funded grain research
is the longest established in Europe and North America,
although similar initiatives were started in the late nineteenth
century in a number of European colonies.
Government-funding bodies have shown a greater interest
in assessing the economic and social impacts of research that
they fund research funding is increasingly viewed as an
investment rather than as grants. The most significant trend
in terms of research intensity is the very significant growth over
the last couple of decades in the level of grain (especially
cereals) R&D carried out by the private sector or alternatively
funded by the private sector but carried out in universities or
government organizations. Most publicly funded agricultural
research in North America and Europe is concentrated in
436 Encyclopedia of Food Grains, Second Edition
the United States, France, and Germany, with Canada and
particularly the United Kingdom having smaller public sector
roles. There is also a greater proportion of agricultural (includ-
ing grains) research carried out in universities in Europe and
North America compared with developing countries, where it
is largely within government-funded research institutes. Grain
R&D carried out by commercial and industry organizations is
reviewed in a separate article, so the emphasis of this article is
on major public sector (government and university) grain
science organizations.
The United States
Although the United States has very large private sector (seed
companies; milling, baking, brewing, and oilseed processing
companies; and biotechnology companies) involvement in
grain R&D, government investments are also significant. A
wide range of temperate grain crops are targeted especially
not only wheat, barley, maize, and oilseeds but also pulses and
subtropical rice, with most resources dedicated to maize and
wheat research. State and federal governments invest roughly
equally in agricultural research.
The Agricultural Research Service (ARS, www.ars.usda.gov)
is the principal research agency of the (national) US Depart-
ment of Agriculture (USDA), and together with the land
grant universities, they form the main research centers that
conduct grain research. The development of a single strategic
plan and national program structure by USDAARS in recent
years has increased the amount of coordination between the
800 or so individual USDA projects underway in different
centers. Crop Production and Protection is the third national
program described in USDAARSs 201217 strategic plan
and delivers science-based information, genetic resources,
and technologies for increased crop productivity, economically
and environmentally sustainable methods of crop production,
and protection from plant diseases and pests. There are six
subprograms, each relevant to grain research: Plant Genetic
Resources, Genomics and Genetic Improvement; Plant Biolog-
ical and Molecular Processes; Plant Diseases; Crop Protection &
Quarantine; Crop Production; and Methyl Bromide
Alternatives.
The two major goals under area 3 are firstly to Protect,
expand, and enhance the United States crop genetic resource
base, increase scientific knowledge of crop genes, genomes,
biological processes and systems, and deliver economically
and environmentally sound technologies that improve the
production efficiency, quality, health and value of the nations
crops and secondly to Develop knowledge, strategies, sys-
tems and technologies that maximize the production efficiency
of our annual, perennial, greenhouse and nursery cropping
systems. Develop new technologies and tools contributing to
improving these systems to meet current and future food crop
http://dx.doi.org/10.1016/B978-0-12-394437-5.00056-5
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
production needs of diversified consumers, while ensuring
economic and environmental sustainability and production
efficiency, health, and value of our nations crops. The Natural
Resources and Sustainable Agricultural Systems program clus-
ter is also relevant.
Some USDA centers are free-standing multidisciplinary
research centers, while smaller ones are typically colocated on
university campuses. Many centers have the mandate to carry
out research directed at the particular agroecological zone in
which the institute is based. For example, wheat is grown in
many parts of the United States, but there are different quality
types suited for different locations soft red winter wheat in
the Midwest, Virginia, and North Carolina; hard white and
some hard red winter wheat in Pacific Northwest; hard red
winter varieties in the Great Plains, from Texas to Montana
on the Canadian border; and spring varieties in northern
central states such as Minnesota, North and South Dakota,
and Montana and appropriately located USDA centers
develop germ plasm and agronomy packages for the different
types of varieties.
Some of the USDA centers carrying out grain research are
based in the following:
Aberdeen, ID Small Grains and Potato Germplasm
Research
Albany, CA (Western Regional Research Center) Genomics
and Gene Discovery and Crop Improvement/utilization
Research
Ames, IA Corn Insects and Crop Genetics Research
Beltsville, MD (Beltsville Agricultural Research Center)
Sustainable Agricultural Systems Laboratory, Instrumentation
and Sensing Laboratory, National Germplasm Resources
Laboratory, Soybean Genomics and Improvement Labora-
tory, and Food Quality Laboratory
College Station, TX (Southern Plains Agricultural Research
Center) Crop Germplasm Research and Insect Control
and Cotton Disease Research
Colombia, MO Plant Genetics Research
Corvallis, OR Forage Seed and Cereal Research
Fargo, ND Cereal Crops Research and Hard Spring and
Durum Wheat Quality Laboratories
Fort Collins, CO National Center for Genetic Resources
Preservation
Ithaca, NY Plant Genetic Resources, Plant Protection
Research, Plant, Soil and Nutrition Research
Lane, OK (South Central Agricultural Research Laboratory)
Genetics and Production Research
Lincoln, NE Grain, Forage, and Bioenergy Research
Lubbock, TX Cropping Systems Research Laboratory and
Plant Stress and Germplasm Development Research
Madison, WI Cereal Crops Research
Manhattan, KS (Center for Grain and Animal Health
Research) Grain Quality and Structure Research, Stored
Product Insect Research, and Hard Winter Wheat Genetics
Research
Peoria, IL (National Center for Agricultural Utilization
Research) Cereal Products and Food Science Research,
Plant Polymer Research, Mycotoxin Research, New Crops
and Processing Research, Crop Bioprotection Research, and
Bacterial Foodborne Pathogens & Mycology Research
437
Pullman, WA (Pacific West Area) Wheat Genetics, Quality
Physiology and Disease Research, Land Management and
Water Conservation Research, and Root Disease and
Biological Control Research
Raleigh, NC Soybean and Nitrogen Fixation Research and
Plant Science Research Laboratories
Shafter, CA Western Integrated Cropping Systems
Research
Sidney, MT Agricultural Systems Research Unit
Stillwater, OK Wheat, Peanut, and Other Field Crops
Research, Plant Science and Water Conservation Research
Stoneville, MS Crop Genetics & Production Research Unit
St Paul, MN Midwest Area Cereal Disease Laboratory and
Plant Science Research
Stuttgart, AR Dale Bumpers National Rice Research Center
Tifton, GA Crop Genetics and Breeding Research and
Crop Protection and Management Research
Urbana, IL Soybean/maize Germplasm, Pathology, and
Genetics Research
West Lafayette, IN Crop Production And Pest Control
Research
Wooster, OH Corn, Soybean and Wheat Quality Research
Wyndmoor, PA Crop Conversion Science & Engineering
Research Unit
USDA also has Economic Research Service (www.ers.usda.
gov), which carries out research and communication activities
in many areas relevant to grains, including farm-level risk
management, commodity outlook research, and economic
and statistical research on marketing and production sustain-
ability. The USDA National Institute of Food and Agriculture
(NIFA, www.nifa.usda.gov) replaced the Cooperative State
Research, Education, and Extension Service in 2009 and is
intended to consolidate all federally funded agricultural
research (including federal support to land grant universities)
and have overarching federal coordination of extension work
done through USDA.
The land grant college system was established in 1862
the name is derived from the fact that the colleges were origi-
nally endowed by grants of public lands in the expanding
western United States. The 1887 Hatch Act established the
agricultural experiment station system to enable these colleges
to undertake research, and the 1914 SmithLever Act estab-
lished the cooperative extension system. This unique system
has the advantage of linking state government R&D and exten-
sion activities to university research. The greatest emphasis for
state government efforts is on breeding new cultivars and
adapting crop production approaches to local conditions and
on education. The universities usually have experiment sta-
tions at different locations in their state, again often corre-
sponding to different agroecological zones but also requiring
researchers to carry out a local outreach function. There are
usually a balance of applied grain research on constraints to
productivity in a particular state and a proportion directed
toward more fundamental research. Cooperative extension
systems from land grant universities have a state development
mandate and often involve location of farm advisors in
counties throughout the state. The continued involvement of
the government and public funding in agricultural extension in
the United States is in contrast with the winding back or exiting
438 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
of state or national government staff from extension roles in
other countries such as the United Kingdom, Australia, and
New Zealand. A summary of some of the main US universities
that carry out grain research is provided in Table 1. Land grant
Table 1 Some US universities undertaking significant grain research
University (website)
Colorado State University, Fort Collins, CO
(www.agsci.colostate.edu)
Cornell University, Ithaca, NY
(www.cals.cornell.edu)
Iowa State University, Ames, IA (www.ag.
iastate.edu)
Kansas State University, Manhattan, KS
(www.k-state.edu)
Michigan State University, East Lansing, MI
(www.maes.msu.edu)
Montana State University, Bozeman, MT
(www.montana.edu/agriculture)
North Carolina State University, Raleigh, NC
(www.cals.ncsu.edu)
North Dakota State University, Fargo, ND
(www.ag.ndsu.nodak.edu)
Oklahoma State University, Stillwater, OK
(www1.dasnr.okstate.edu)
Ohio State University, Columbus, OH
(www.oardc.ohio-state.edu)
Pennsylvania State University, University
Park, PA (www.research.cas.pse.edu)
Purdue University, West Lafayette, IN
(www.agriculture.purdue.edu/arp)
Rutgers University, New Brunswick, NJ
(www.foodsci.rutgers.edu/caft)
NJ Agricultural Experiment Station
(www.cook.rutgers.edu)
Texas A&M University, College Station, TX
(www.agresearch.tamu.edu; www.
agrprogram.tamu.edu)
University of Arkansas, Fayetteville, AR
(www.uark.edu/admin/aes)
University of California, Davis (www.ucanr.
org; www.caes.ucdavis.edu)
University of Illinois, UrbanaChampaign, IL
(www.web.aces.uiuc.edu)
University of Idaho, Aberdeen, ID (www.
ag.uidaho.edu)
University of Minnesota, St. Paul, MN
(www.maes.umn.edu)
University of Missouri, Colombia, MO
(www.cafnr.missouri.edu)
University of Nebraska, Lincoln, NE
(www.unl.edu)
Washington State University, Pullman, WA
(www.cahe.wsu.edu)
universities are usually predominantly state-funded, but with
significant federal funding as well.
In 2011, the US government commenced the presidents
global hunger and food security initiative (www.feedthefuture.
Grain research emphasis
Genomics, biotechnology, dryland farming systems, wheat breeding, precision agriculture
Genetics, biotechnology, basic plant science, maize breeding, seed science, cereal pathology, cereal
product rheology. Wheat variety development, genetics research, disease resistance, and crop
management. Particular emphasis on breeding methodologies, disease resistance, comparative
genomics, association mapping, allele characterization, and genomic selection methods
Corn and soybean breeding, nonfood crop products, corn quality, basic plant sciences, crop genomics,
seed technology
Wheat, soybean, and sorghum breeding; cereal (wheat, corn, sorghum, and millet) agronomy and
physiology, entomology, cereal chemistry, milling and baking, livestock feeds. Wheat genetics,
development of genomics resources and high throughput genotyping technologies, wheat breeding,
germ plasm development
Grain crop breeding, cereal chemistry, genomics
Winter wheat, barley, durum, pulses, canola, and soybean breeding, precision agriculture, integrated
pest management
Bioinformatics, integrated pest management
Basic research on wheat quality and processing; milling; pasta production; brewing; farming systems,
pathology, and entomology
Baking, grain protein chemistry, oilseed chemistry, plant diseases
Maize production systems, cereal and oilseed quality, soft red winter wheat and soybean agronomy and
breeding, soy protein functionality
Starch chemistry, entomology
Plant stress physiology, biotechnology, integrated pest management, precision agriculture, new grain
products, carbohydrate research
Cereal processing, plant biotechnology
Precision agriculture, farming systems, plant genomics, grain food processing
Rice, wheat, oat, and soybean breeding; agronomy and physiology, rice processing
Genetic resources conservation, genetics of cereals and legumes, agronomy, wheat breeding,
entomology, plant pathology, rice breeding
Breeding, Marker-Assisted selection, Positional cloning of agronomic genes, Dissection of flowering
and disease resistance regulatory pathways
Maize and soybean breeding and agronomy, cereal diseases, starch chemistry
Grain breeding and agronomy (wheat, oilseeds)
Cereal structure, wheat and barley breeding and pathology, soybean diseases, precision agriculture,
cereal food processing
Tillage, soybean, sorghum, and corn cropping systems and soybean breeding, basic grain science,
maize genome mapping
Processing of grain starches, cereal breeding, genomics. Wheat breeding and genetics of winter hardy,
disease and insect resistant, high-yielding cultivars. Fundamental research in wheat virology, fungal
disease control, biochemistry, statistical approaches, and breeding methodology including
transformation.
Grain crops breeding and genetics, pathology/entomology, soil science, processing
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
gov/approach), which includes the Feed the Future research
strategy. This is a major international development assistance
initiative managed through USAID. Investments under this
strategy range from longer-term research to address major
global challenges to applied and adaptive research guided by
host-country priorities for nearer-term impact. A number of US
research laboratories within USDA and the university sector are
involved given the central role of grains in food security, it is
not surprising that several initiatives support grain research.
Within three broad research themes (advancing productivity,
transforming production systems, and enhancing food safety
and nutrition), these include work on wheat stem rust, abiotic
stress tolerance, genetic resources, marker-assisted technolo-
gies, legume productivity, and management of mycotoxin con-
tamination in stored grains.
Canada
The majority of wheat breeding and research in Canada is
conducted by the public sector, although private sector breed-
ing is gaining momentum and Syngenta has released wheat
varieties in Canada since 1993. Several publicprivate partner-
ships exist between industry and most publically funded
research institutions. Agriculture and Agri-Food Canada
(www.agr.gc.ca/science and innovation/research centers)
carries out grain research at several of its 19 research centers
across the country. Cereal research is carried out mainly at
the Winnipeg (Manitoba) Cereal Research Centre, which is co-
located with the University of Manitoba and emphasizes six
areas: cereal diseases, cereal germ plasm and genomics, pulse
germ plasm, nutrition and functional foods, bioprospecting
from grains, and grain storage research. The Eastern Cereal
and Oilseed Research Centre (Ottawa, ON) develops new vari-
eties of spring and winter wheat, barley, corn, oats, and soy-
beans and has the central biodiversity mandate for Canada.
There is a strong emphasis on gene isolation, gene transfer,
and the study of gene expression in grain crops, bioprospecting
for new bioactives, and study of Fusarium resistance.
Wheat breeding is also carried out at the Semiarid Prairie
Agricultural Research Centre (Swift Current, Saskatchewan,
along with breeding of other cereals) and the Lethbridge
Research Centre (Alberta). The Lethbridge Research Centre
also carries out bean and forage crop breeding and research
on water, soil, crop, and manure management for prairie con-
ditions. There is an emphasis on disease and disease resistance
and agronomy. The Brandon Research Centre (Manitoba) is
the main site of barley breeding along with a strong soil science
focus, while the Lacombe Research Centre (Alberta) under-
takes oat-varietal improvement research, along with barley
breeding (disease resistance focus) and farming systems
research. The Saskatoon Research Centre (Saskatchewan) is
the main center of breeding programs for canola and mustard
for prairie conditions. Some canola breeding and breeding for
other short-season grain crops also take place at the Lacombe
Research Centre, while the Greenhouse and Processing Crops
Research Centre in Harrow, Ontario, also carries out soybean
breeding. Agronomy research for grain production is under-
taken at the Soils and Crops Research and Development
439
Centre, Quebec (for cool, wet production systems), Atlantic
Cool Climate Crop Research Centre (St Johns, Newfound-
land), Crops and Livestock Research Centre (Charlottetown,
Prince Edward Island), Brandon Research Centre, and
Semiarid Prairie Agricultural Research Centre (Swift Current,
Saskatchewan).
Several universities carry out grain research in Canada,
often working closely with Agriculture and Agri-Food Canada
scientists. These include the following:
University of Saskatchewan (www.usask.ca) with research
emphasis on wheat, barley, oat, canola, and legume (lentil,
pea, dry bean, and chickpea) breeding and agronomy, plant
genetic engineering, molecular marker development, and
malting and brewing science.
University of Alberta, Edmonton (www.afns.ualberta.ca),
studies grain and oilseed crops for Western Canada
breeding, disease resistance, agronomy, and livestock feed
utilization. In wheat, it focuses on conventional breeding
and genetic research with an emphasis on adaptation of
wheat to the short growing season on the Canadian prairies
and agronomy and breeding of grains for organic agricul-
tural environments.
University of Manitoba, Winnipeg (www.umanitoba.ca/
afs/plant_science), carries out research on canola and
wheat breeding and postharvest technology, grain storage
technology, cereal chemistry, baking technology, pulse
food science, functional foods, and cereal agronomy
including precision agriculture, genetics and wheat breed-
ing, agronomic production systems, pathology, breeding
methodologies, and end-use quality.
University of Guelph, Ridgetown, Ontario (www.ridge-
townc.com/index.cfm), emphasizes grain crops breeding
(soybean, barley, wheat, and canola), agronomy, pathol-
ogy, and genome mapping.
McGill University, Quebec, Ontario (www.mcgill.ca), is
involved with basic plant science, biotechnology, and
some grain food processing research.
The National Research Council (NRC) Plant Biotechnology
Institute in Saskatoon (www.pbi.nrc.ca) carries out basic
research in genomics and proteomics and has a special inter-
est in the modification of oil composition and of agronomic
traits of oilseeds and in cereal and legume molecular marker
and transformation technology. Some individual provinces
also have agencies that carry out applied agriculture and
food research. For example, the Field Crop Development
Centre is funded in part by the province of Alberta and
conducts breeding and development programs focussed on
feed and malting barley, spring and winter triticale, and
winter wheat. The Canadian Grains Commission in Winni-
peg, Manitoba, regulates grain handling and establishes and
maintains grain quality standards for Canada. Its Grain
Research Laboratory (www.grainscanada.gc.ca/cgc-ccg/grl-lrg/
grlm-mlrg-eng.htm) has emphasis on research to understand-
ing the genetic, environmental, structural, and biochemical
basis on end-use quality of major grains, how grading factors
affect end-use qualities, and investigating new uses for Cana-
dian grain. The Canadian International Grains Institute
(www.cigi.ca), also in Winnipeg, has a technical training
and information dissemination role.
440 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
Mexico
In Mexico, maize is the main grain crop, but the national
program played an important role in semidwarf wheat breed-
ing ahead of the establishment of Centro Internacional de
Mejoramiento de Maiz y Trigo, International Wheat and
Maize Improvement Center (CIMMYT). Most federal govern-
ment agricultural research is carried out by the Instituto Nacio-
nal de Investigaciones Forestales Agrcolas y Pecuarias (INIFAP,
www.inifap.gob.mx); some of the state governments have
small grain research activities. There is also significant activity
within the higher education and science and technology min-
istries Consejo Nacional de Ciencia y Tecnologa (CONACYT,
www.conacyt.mx), with grain research being carried out in
several of their regional centers (Centro de Investigacion en
Alimentacion y Desarrollo (CIAD), AC, Centro de Investigaci-
on y Assistencia en Tecnologa y Diuseno del Estado de Jalisco
(CIATEJ), Centro de Investigaciones Biologicas del Noroeste
(CIBNOR), and El Colegio de la Frontera Sur (ECO SUR)).
Postgraduate colleges, within the universities managed under
the agriculture secretariat (SAGAR), along with the agronomy
faculties in autonomous universities in different Mexican
states, are also active. Mexico is home to a number of national
seed companies (maize, sorghum, and wheat), along with sub-
sidiaries of foreign (especially the United States) seed
companies, and these carry out a variety of trials and agronomy
work.
Europe
The Seventh Framework Programme for Research (www.ec.
europa.eu/research/fp7/index_en.cfm) provided 50 billion
for collaborative research from 2007 to 2013 across ten the-
matic areas, one of which is bioeconomy, encompassing food,
agriculture and fisheries, and biotechnology (www.ec.europa.
eu/research/bioeconomy/index_en.htm). Grain research fea-
tures in several projects and programs with the bioeconomy
area, including improved feed utilization of grain crops, crop-
ping system resilience under climate change, value addition to
crops through biotechnology and food quality and safety, and
a range of major basic plant science and biotechnology
projects.
In Western Europe, yields of most major cereals are high, and
it is clear that there has been a shift away from research on
increasing production to an emphasis on environmentally
friendly production and managing variability under climate
change. While biotechnology programs are strong, there is lim-
ited work on crop genetic transformation due to lack of accep-
tance of commercial genetically modified (GM) food crops by
many European governments. Other areas of grain research
emphasis across Europe include quality and processing technol-
ogies, including value-addition and nonfood uses of grains; food
safety, nutrition, and health (with grains, the main emphasis is
on mycotoxins, nutraceuticals, dietary fiber, and allergies and
intolerances); traceability (e.g., of GM cereals); and breeding,
genomics, and diagnostic technologies.
Among the UK government institutes, the Biotechnology
and Biological Sciences Research Council (BBSRC, www.bbsrc.
ac.uk) emphasizes research on grains within several institutes:
John Innes Centre, Norwich prebreeding and basic research
on breeding technology for yield and quality, plant biotech-
nology, and biology of cereals and brassica crops
Rothamsted Research grain crop performance, farming
systems, seed science, GM wheat trialing, and plant
bioscience
Institute of Biological, Environmental and Rural Sciences
(at Aberystwyth University) genome diversity analysis,
breeding methodologies, oat breeding, and biofuels
The Genome Analysis Centre
The James Hutton Institute in Scotland (www.hutton.ac.uk)
conducts research on the genetics of quality relationships in
barley and wheat, farming systems, weeds, pests, and diseases.
Major British universities carrying out research in grain science
include the University of Reading (www.apd.rdg.ac.uk); Univer-
sity of Nottingham, Sutton Bonington (www.nottingham.ac.
uk); University of Bristol, Long Ashton; University of East Anglia,
Norwich (www.uea.ac.uk); Heriot-Watt University, Edinburgh;
University of Plymouth, Devon; University of Leeds; University
of Manchester; Institute of Science and Technology; Satake Cen-
tre for Grain Process Engineering (www.ce.umist.ac.uk); and
Queens University BelfastSchool of Agriculture and Food Sci-
ence (www.qub.ac.uk). The Home-Grown Cereals Authority
(www.hgca.com) is an organization funded on grower levies to
support research and provide market information and extension
information for British grain farmers. Campden BRI (www.
campdenbri.co.uk) provides technical and research support for
industry and incorporates three former centers the Chorley-
wood Food Research Association, Brewing Research
International, and Campden Food Research Institute.
In Ireland, the Irish Agriculture and Food Development
Authoritys (TEAGASC, www.agresearch.teagasc.ie) Oak Park
Crops Research Centre emphasizes research on grain crops,
especially aiming for more efficient production systems for
cereals, grain lupins and rape oilseed. There are a strong
emphasis on the management of fungal and viral diseases of
cereal crops and an increasing focus on nonfood uses for grain
crops. The main university for production research on grain
crops is University College Dublin, while University College
Cork emphasizes grain processing research. In Switzerland, the
major government grain research institutes are the Federal
Office for Agriculture, Swiss Federal Research Station for
Plant Production, and Swiss Federal Research Station for Agro-
ecology and Agriculture, while in the university sector, the
Swiss Federal Institute of Technology in Zurich carries out
most of the applied grain sciences research.
In France, the Institut National de la Recherche Agronomique
(INRA, www.inra.fr/en) is the main government research orga-
nization in agriculture and grain research within four divisions:
Genetics and Plant Breeding, Plant Health and Environment,
Environment and Agronomy, and Science and Process Engineer-
ing of Agricultural Products. Major areas of grain research at
INRA are structural and functional genomics, analysis of genetic
diversity, plant breeding, hostpathogen interactions, and grain
development and quality. INRA has a large center in Nantes
with a research emphasis on agriculture and food processing.
There is an additional emphasis on cereal food and nutritional
quality and nonfood uses, with a strong emphasis on processing
technologies and basic protein and starch molecular sciences
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
related to cereals. At INRA Montpellier, there is an ongoing
research on processing technology, grain crop production
systems, plant protection, and genetic resource conservation
as well as more basic studies on grain crop development.
Several French universities have Ecole Nationale Superieure
Agronomique, for example, in Paris Grignon, (the Institut
National Agronomique (INA-PG)), Montpellier, Rennes,
Nancy, and Toulouse. ARVALIS Institut du vegetal (www.arva-
lisinstitutduvegetal.fr/en) is responsible for applied research
on cereals, maize, pulses, potato, and forage crops. The activ-
ities of this technical institute, which is run and financed by
farmers, include sustainable cropping systems, precision farm-
ing, remote sensing, satellite images, biotechnology, farm eco-
nomics, quality assurance, safety quality, traceability, human
food, animal feed, and technology transfer. CETIOM (www.
cetiom.fr) is responsible for similar topics on oil crops. Cer-
eales Vallee (www.cereales-vallee.org) is a government
competitiveness cluster established in 2005. It supports the
development of applied cereal research projects through col-
laboration between a number of public and private partners in
France. There are four programs that in turn address sustain-
able cereal production, cereals for human nutrition and health,
cereals for animal feed, and cereals for agromaterials.
Breeding of new cereal cultivars in Germany is aimed at
increased yields, improved resistances to biotic and abiotic
stress, and high quality and like many other European coun-
tries is now conducted by private breeding companies, with
government researchers carrying out prebreeding activities.
Cereal research is conducted at the following:
Universities, for example, the Technical University of
Munich, Technical University of Berlin, Martin Luther Uni-
versity Halle-Wittenberg, Justus Liebig University Giessen,
Humboldt State University, and the Universities of Bonn,
Gottingen, Hohenheim, Kassel, Kiel, Rostock, and
Paderborn.
Federal research institutes, for example, the Leibniz Insti-
tute of Plant Genetics and Crop Plant Research (IPK, www.
ipk-gatersleben.de/en); the Julius Kuhn Institute (JKI, www.
jki.bund.de/en); Federal Centre for Cereal, Potato, and
Lipid Research (BAGKF, Detmold, www.mri.bund.de/en/
de/max-rubner-institut/geschichte/bundesanstalt-fuer-
getreide-kartoffel-und-fettforschung.html); the Federal
Research Centre for Plant Breeding (BAZ, www.genres.de);
and the Institute of Agricultural Engineering (ATB,
Potsdam-Bornim, www.atb-potsdam.de/en.html).
Max Planck Institute for Plant Breeding Research (MPIPZ,
www.mpipz.mpg.de/2169/en).
German state research centers in agriculture several of
these carry out grain research, sometimes in conjunction
with universities (e.g., Bavarian State Research Centre for
Agriculture and State Plant Breeding Institute of the Univer-
sity of Hohenheim (Baden-Wurttemberg)).
The proWeizen initiative was recently founded and is coordi-
nated by the German Federation of Private Plant Breeders (GFP).
Its aim is to coordinate and foster German wheat research. It
consists of leading research institutes and private companies and
organizations involved in wheat breeding and growing.
In the Netherlands, two institutes within the Dienst Land-
bouwkundig Onderzoek (DLO), a private nonprofit research
441
institute that is now part of Wageningen University and Research
Center, carry out grain research, including the Agrotechnology
and Food Sciences Group and Plant Research International
(www.pri.wur.nl/uk) who focus on crop protection, crop ecol-
ogy, agricultural systems, genetics, and reproduction. The Neth-
erlands Organisation for Applied Scientific Research (TNO)
conducts research on cereal processing. Wageningen University
(www.wageningenur.nl/en/wageningen-university.htm) is the
main university research center for grain research. In Belgium,
there are relevant government agricultural research centers
in Gent and Gembloux, while at least four universities carry
out some aspects of grain production and processing research
Ghent University, Catholic University of Leuven, Gembloux
Agricultural University, and Catholic University of Louvain.
The Italian government Ministry of Agricultural, Food and
Forestry Policies has several research institutes covering grain
research, including various Research Institutes for Cereal
Research (ISC, www.sito.entecra.it/portale/cra_dati_istituto.
php?id205&linguaEN) and the Industrial Crops, Plant
Nutrition, Plant Pathology and Food and Nutrition
(IMRAN). The NRC (CNR, www.cnr.it) has a separate system
of research institutes, and some of these also carry out grain
research. Many major states in Italy host universities that have
agriculture faculties, and most of these carry out at least some
grain research the Universities of Turin, Milan, Padua, Udine,
Bologna, Piacenza, Florence, Pisa, Ascoli, Perugia, Viterbo,
Naples, Bari, Potenza, Reggio Calabria, Catania, Palermo, and
Sassari.
The Cereal Institute, National Agricultural Research Foun-
dation (NAGREF, www.nagref.gr), Thessaloniki, Greece, is the
Greek government research institute responsible for cereal
breeding and cereal food product development. However, the
Ministry of Agriculture has a separate research service with a
network of institutes in different parts of the country. At
least three universities carry out some grain research work
Agricultural University of Athens, Aristotle University of
Thessaloniki, and the Mediterranean Agronomic Institute at
Chania. In Hungary, the Ministry of Agriculture and Rural
Development manages the Central Food Research Institute
(KEKI) and the Agricultural Biotechnology Center (MBK), as
well as the Cereal Research Institute in Szeged. The Hungarian
Academy of Sciences has several institutes relevant to grain
science the Agricultural Research Institute, the Research Insti-
tute for Soil Science and Agricultural Chemistry, and the Plant
Protection Institute. Universities with research areas most rel-
evant to grain science are the Technical University of Budapest
(grain processing) and the Universities of Agricultural Sciences
at Godollo, Debrecen, and Pannon.
In Austria, the Federal Ministry of Agriculture, Forestry,
Environment and Water Management has research institutes
for general agriculture and an institute of cereal processing,
applied soil science, and agricultural engineering. There is also
the University of Natural Resources and Life Sciences in Vienna.
The International Association for Cereal Science and Technol-
ogy (www.icc.or.at) is headquartered in Vienna and has
evolved from being a professional organization of cereal scien-
tists to also hosting a number of EU (Seventh Framework)-
funded grain research projects. These include projects on
food-related disease prevention, bioactives and food supple-
ments from cereal grains, food allergens and allergy risk
442 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
management, and mycotoxin analysis. In the Czech Republic,
relevant Ministry of Agriculture research institutes include the
Food Research Institute, Research Institute of Crop Production,
Research Institute for Soil and Water Conservation, and
Research Institute of Agricultural Engineering. There are semi-
commercial institutes responsible for cereals (Agricultural
Research Institute Kromeriz Ltd.), oilseed crops (Oseva Ltd.),
and brewing and malting. Two universities Mendel University
of Agriculture and Forestry and Czech University of Agriculture
are relevant to grain research. In Spain, the Ministry of Agri-
culture, Fisheries and Food runs the National Institute for
Agriculture and Food Research and Technology (INIA, www.
inia.es), while the Polytechnic University of Madrid has a large
faculty involved with research in agriculture. There is also an
INIA in Portugal, while agricultural research is carried out at the
Technical University of Lisbon, the Universidade de Tras-os-
Montes e Alto Douro, and Universidade dos Acores.
In Sweden, the government Swedish Institute for Food
Research and the Swedish Institute for Food and Biotechnol-
ogy (SIK, a joint industry/government body) carry out grain
processing research. Production research and basic scientific
investigations within universities include the Swedish Univer-
sity of Agricultural Sciences at Svalov and Lund University.
Three main institutes in Norway carry out grain research.
Within the government Ministry of Agriculture, there is the
State Agricultural Research Station (SFL) and the Agricultural
University of Norway at Aas, while the Norwegian Food
Research Institute (Matforsk) is a nonprofit organization. In
Finland, the Ministry of Agriculture and Forestry has the Agri-
cultural Research Centre of Finland and Plant Breeding Insti-
tute, while VTT Biotechnology in Espoo carries out cereal
technology research in close collaboration with industry. The
University of Helsinki is the main higher education center
dealing with grain research. In Denmark, the Ministry of
Food, Agriculture and Fisheries has Institutes of Agricultural
Sciences and Agricultural Engineering. The Royal Veterinary
and Agricultural University also carries out grain research.
Activities related to the Carlsberg laboratory are described in
the article Research Organizations of the World: Global
Trends and the Commercial Sector.
In Russia, the Russian Academy of Agricultural Sciences
(www.euracadagri.com/eng/members/russia.php) comprises
many dozen institutes. These include All-Russia Research Insti-
tutes (e.g., All-Russia Research Institute of Plant Quarantine
(Bykovo), Plant Protection (St. Petersburg), and Biological
Protection of Plants (Ekaterinburg), which carryout entomol-
ogy research related to grains) and regional research institutes
in former republics. The latter often cover a range of agricul-
tural disciplines and some have an almost 100-year history;
many have significant grain breeding and seed distribution
programs. Three of the All-Russia Research Institutes most
relevant to grains are the Institute of Legumes and Groat
Crops (Streletskoye), Institute of Sorghum and Cereals (Zerno-
grad), and N.I. Vavilov Institute of Plant Industry (www.vir.
nw.ru). The latter institute, based in St. Petersburg, but with a
number of experimental stations throughout Russia, is the
main internationally renowned Russian institute emphasizing
the conservation and study of plant genetic resources, includ-
ing grain crops. Most states have state agrarian universities, and
there are a small number of specialist agrarian academies
including Moscow Agricultural Academy.
After separation from Russia in 1991, national agricultural
research systems developed in most of the former Soviet
republics using the institutional assets left by the former
USSR; several of these have a crop emphasis although resources
for research are very limited and it has been difficult (and often
undesirable) to maintain the staff numbers and infrastructure
left behind. They have had to develop their priority setting to
match specific requirements of their various agroecological
zones and the changed policy/economic environment includ-
ing a greater emphasis on markets rather than a central procure-
ment and distribution system.
Exercises for Revision
What are some of the major institutional trends in grain
research in North America and Europe over the last
decade?
How do the major grain research priorities differ between
now and 30 years ago?
What are the major priorities globally for biofuel research?
Name three of the countries within Europe and North
America that have the highest proportion of publicly
funded agricultural R&D.
Why are agricultural universities in the United States called
land grant colleges?
Exercises for Readers to Explore the Topic Further
Explore the major grain research projects funded under the
Seventh Framework Programme of the European Union.
What are the common themes?
What have been the major achievements of the N.I. Vavi-
lov Institute of Plant Industry in Russia over the last
century?
What areas of grain research from the Netherlands major
agricultural university in Wageningen have had global
impact?
See also: Agronomy of Grain Growing: Plants: Diseases and
Pests; Soybean: Agronomy; Wheat: Agronomy; Breeding of Grains:
Barley: Genetics and Breeding; Canola/Rapeseed: Genetics and
Breeding; Maize: Breeding; Soybean: Germplasm, Breeding, and
Genetics; Wheat Breeding: Exploiting and Fixing Genetic Variation by
Selection and Evaluation; Genetics of Grains: Development of
Genetically Modified Grains; Genome Mapping; Genomics of Food
Grains; Grain Marketing and Grading: Barley: Grading and
Marketing; Wheat: Grading and Segregation; Wheat: Marketing;
Grains Around the World: Grain Production and Consumption:
Cereal Grains in North America; Grain Production and Consumption:
Europe; Grain Production and Consumption: Oilseeds in North
America; Grain Production and Consumption: Overview; Non-food
Products from Grains: Biodiesel; Cereal Grains as Animal Feed; Fuel
Alcohol Production; Scientific Organizations Related to Grains:
Scientific Societies Associated with Grain Science; The Basics: The
Grain Crops: An Overview; The Cereal Grains: An Overview of the
Family of Cereal Grains Prominent in World Agriculture.
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organizations of the World: Europe and North America
Further Reading
Fox JA and Ward LL (2016) Grain production and consumption: Cereal
grains in North America. In: Wrigley C, Seetharaman K, Corke H, and
Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1, pp. 391400.
Elsevier Ltd.
Grassini P, Estraidge KM, and Cassman KG (2013) Distinguishing between yield
advances and yield plateaus in historical crop production trends. Nat. Commun.
4: 2918. http://dx.doi.org/10.1038/ncomms3918.
Lindhaeur MG (2016) Grain production and consumption: Europe. In: Wrigley C,
Seetharaman K, Corke H, and Faubion J (eds.) Encyclopedia of Food Grains,
2nd edn., vol. 1, pp. 383390. Elsevier Ltd.
443
Pardey RG and Beintema NM (2001) Slow Magic: Agricultural R&D a Century after
Mendel. Washington DC: International Food Policy Research Institute.
McVetty PBE, Lukow OM, Hall LM, Rajcan I, and Rahman H (2016) Grain production
and consumption: Oilseeds in North America. In: Wrigley C, Seetharaman K,
Corke H, and Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1,
pp. 401408. Elsevier Ltd.
Ray DK, Mueller ND, West PC, and Foley JA (2013) Yield trends are insufficient to
double global crop production by 2050. PLoS One 8(6). http://dx.doi.org/10.1371/
journal.pone.0066428.
Skerritt JH (2016) Research Organizations of the World: CGIAR. In: Corke H, Wrigley C,
Faubion J, and Seetharaman K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1,
pp. 429435. Elsevier Ltd.
 Research Organizations of the World: Asia-Pacific, CentralSouth America, and
AfricaMiddle East
JH Skerritt, University of Queensland, St Lucia, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Countries in regions other than North America and Europe
vary significantly in their grains research capacities.
Within these regions, Australia, Brazil, China, and India
have the greatest grains research capacity.
Global grain price spikes in the last few years have increased
the emphasis on grains research for food security in devel-
oping countries.
Learning Objectives
To identify and understand the major priorities of the
major organizations in grains research in countries other
than North America and Europe
Introduction
Many of the trends in North American and European grains
research and grains research organizations are reflected in the
developed and major developing economies within the Asia-
Pacific, Latin American, African, and Middle Eastern regions, but
to varying degrees. In Australia and India, there has been a move
toward formation of networks between research institutes, often
being a requirement of funding program design. In Australia,
China, and Brazil, the private sector has significantly increased
its role, particularly in breeding. Other trends in Australia
include the greater importance of levies from grain growers in
supporting research in a number of countries and the increase in
commercial involvement in grains breeding and biotechnology.
Across each of the regions reviewed in this article, a shift in
focus of many of the grain R&D institutions took place in the
1990s and first few years of the twenty-first century from a
largely production focus to an emphasis on environmental
issues such as water-use efficiency and sustainability of farming
systems. However, with grain price spikes that occurred over the
last few years, several Asian countries have reintroduced a food
security imperative to the top of their national agricultural R&D
agenda. Within Asia, exports of grains such as rice are significant
contributors to national incomes for countries such as Thailand
and Vietnam, while grain import requirements have critical
balance of payments and thus political imperatives for India,
China, Indonesia, and the Philippines. While significant excep-
tions exist, an institutional challenge for developing countries is
the typically inadequate levels of collaboration between research
institutes within and between countries especially when
comparing the performance of different lines in different agro-
ecological zones.
For the developing countries that benefited from the
higher-yielding varieties of the green revolution, sustaining
the productivity gains through more efficient use of water and
444 Encyclopedia of Food Grains, Second Edition
prevention of erosion, waterlogging, salinity, and acidification
is now critical. Also critical is research on grain quality and
addressing market requirements, as grain production in several
of these countries struggles to compete with subsidized Euro-
pean and US agriculture.
Major Public Sector (Government and University)
Grain Science Organizations
Australia and New Zealand
There are three main public sector providers in grains research
in Australia state government departments, Commonwealth
Scientific and Industrial Research Organisation (CSIRO), and
universities.
Unlike in many countries, the national Department of Agri-
culture (www.agriculture.gov.au) does not conduct laboratory
or field research in agriculture.
In the last 1015 years, there has been a shift in emphasis of
many state departments from an overwhelming emphasis on
production agriculture to sustainable production, and over the
last decade, as budgets become more constrained, a decrease in
research resources. In this period, grains breeding in Australia,
particularly of wheat, has moved into the private sector, with
public research agencies instead carrying out germplasm devel-
opment and prebreeding activities. While one company (Long-
Reach Plant Breeders) is freestanding, the three others
(Australian Grain Technologies, InterGrain, and HRZ
Wheats) are consortia involving private industry and state or
national research agencies. Some of the states retain strong
capacity in grains prebreeding and agronomy and in soil sci-
ence and farming systems research. Information on their
research programs is available as follows:
NSW Department of Primary Industries (www.agric.nsw.
gov.au)
Department of Economic Development, Jobs, Transport
and Resources (www.economicdevelopment.vic.gov.au)
Queensland Department of Agriculture, Fisheries and For-
estry (www.dpi.qld.gov.au)
South Australian Research and Development Institute
(SARDI) (www.sardi.sa.gov.au)
Department of Agriculture and Food, Western Australia
(www.agric.wa.gov.au)
Department of Primary Industries, Parks, Water and Envi-
ronment, Tasmania (www.dpiwe.tas.gov.au)
Department of Primary Industry and Fisheries, Northern
Territory (www.nt.gov.au/d/Primary_Industry)
All state governments have formed various formal agricultural
research alliances with universities some include joint
appointment of staff and a single organizational structure,
while others mainly are based on colocation with universities.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00057-7
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World
In some cases, the alliances cover most areas of grains research; in
others, specialized areas such as biotechnology. Examples rele-
vant to grains research include the Queensland Alliance for Agri-
culture and Food Innovation (QAAFI, involving the University of
Queensland, www.qaafi.uq.edu.au); the EH Graham Centre for
Agricultural Innovation (involving Charles Sturt University,
www.csu.edu.au/research/grahamcentre); the Primary Industries
Innovation Centre in NSW (involving the University of New
England, www.dpi.nsw.gov.au/research/alliances/primary-indus-
tries-innovation-centre); AgriBio in Victoria (involving La Trobe
University, www.latrobe.edu.au/agribio); the South Australia
Research and Development Institute, colocated with the Univer-
sity of Adelaide (www.sardi.sa.gov.au); the Western Australian
State Agricultural Biotechnology Centre, located at Murdoch
University (www.sabc.murdoch.edu.au); and the Tasmanian
Institute of Agriculture (www.tia.tas.edu.au). While Queensland
and Victoria retain reasonable capacity, an increasing proportion
of extension services, formerly provided by the state governments,
are now provided to farmers on a commercial basis by the private
sector and/or incorporated farmers (growers) groups.
Within the national research organization CSIRO (www.
csiro.au), much research (including grains research) is now
being conducted in cross divisional flagship programs. The
Food Futures Flagship is using advanced genetics to create dif-
ferentiated grain, food, and feed products, focussing on wheat
barley with improved processing quality and oilseeds with
enhanced oil content and improved nutritional value. Other
flagships carrying out grains research include biosecurity,
sustainable agriculture (natural resource management and farm-
ing systems focus), and climate adaptation flagships. A small
amount of grains-related research is conducted outside flag-
ships, by the Divisions of Plant Industry; Animal, Food and
Health Sciences; Ecosystem Sciences; and Land and Water.
Some independent nongovernment grains research organi-
zations exist including GrainGrowers (www.graingrowers.
com.au), their expertise including milling, near-infrared reflec-
tance technology, baking science and technology, and Asian
wheat foods. The Australian Grains Centre (www.cbh.com.au/
our-business/operations/grain-quality/australian-grain-centre.
aspx) and Agrifood Technology (www.agrifood.com.au)
emphasize testing services.
Relevant grains research is carried out at several universities.
This includes the following:
University of Adelaide bread and durum wheat and barley
prebreeding, molecular markers, cereal root diseases, cereal
pathology, biotechnology, weed management, malting,
and brewing biochemistry
University of Sydney prebreeding for rust resistance in
barley and wheat, cereal diseases, cereal genomics, preci-
sion agriculture, soil science, and crop modeling
University of Melbourne research to underpin the sustain-
ability of temperate grain crop production, with emphasis
on agronomy and farming systems, and protection of nat-
ural resources
University of New England agronomy and soil science,
plant nutrition, crop protection, weed science, and inte-
grated pest management
University of Queensland production systems and post-
harvest handling of cereals, grain legumes and oilseeds, weed
management, plant protection, and molecular biology
445
University of Western Australia grain crop science and
molecular genetics
Charles Sturt University agronomy, irrigation, and plant
pathology
Curtin University agronomy, precision agriculture, and
grains processing
Murdoch University molecular markers for barley and
wheat, grain legume genome mapping, transgenics,
diseases, and viruses
University of New South Wales grain storage engineering
University of South Australia agricultural machinery
research and design
University of Southern Queensland plantpathogen
interactions, cereal genetic engineering, marker-assisted
breeding, and canola breeding
University of Tasmania cereals, grain legumes, and oil-
seeds as break and rotation crops
Over the last 20 years, there has been a significantly increased
emphasis on establishment of more formal links between
research organizations. This has been driven to a significant
extent by the advent of the Cooperative Research Centres
(CRCs, www.crc.gov.au) program and encouragement by fun-
ders. At present (January 2014), there are only two CRCs that
have relevance to grains research (Future Farm Industries CRC
and the Plant Biosecurity CRC) many fewer than earlier in
the program. The Grains Research & Development Corpora-
tion (www.grdc.com.au) facilitates and manages funding for
research and assists in communicating results. Government
and industry R&D levies co-fund this research.
The main New Zealand organization undertaking grains
research is Plant & Food Research (www.plantandfood.co.nz),
one of several government-owned Crown Research Institutes.
Its program includes wheat and barley genomics, breeding and
production systems, and food processing.
Universities with research interests in grains include Massey
University (Palmerston North, www.massey.ac.nz) with a
research emphasis on food science and engineering. Lincoln
University (Christchurch, www.lincoln.ac.nz) carried out
research on grain foods composition, crop agronomy, seed
science and technology, and some biotechnology. Although
wheat is one of the major cereals, the growing environment is
very different from that in Australia, requiring rather different
varietal characteristics. Further information on grain produc-
tion and consumption in Oceania is discussed elsewhere in
this Encyclopedia.
North Asia
In Japan, much of the grains research is carried out in institutes
managed under the Agriculture, Forestry, and Fisheries
Research Council of the Ministry. National Agriculture and
Food Research Organizations (www.naro.affrc.go.jp/english/
index.html) Institute of Crop Science in Tsukuba conducts
research on the breeding of rice, wheat, barley, soybean,
sweet potato, and industrial crops such as sesame and on the
development of new technologies for breeding. Research activ-
ities also cover the understanding of basic mechanism of agro-
nomic and physiological traits regarding crop quality,
photosynthesis, and productivity. The NARO Agricultural
446 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World
Research Center, also headquartered in Tsukuba, focuses on
rice and soybean systems, agronomy, soil fertility, crop nutri-
tion and rotations, pest management, and biofuels. There are
four main regional research centers. The National Institute of
Agrobiological Sciences (www.nias.affrc.go.jp/index_e.html,
also headquartered in Tsukuba) is the largest agricultural
research institute in Japan for basic life sciences, and a number
of its programs focus on grain crop/genomics and genetic
research. Major universities with agriculture faculties include
Tokyo University of Agriculture and Technology, Tohoku Uni-
versity, Kyoto University, Kyushu University, Hokkaido Uni-
versity, Kagoshima University, Yamaguchi University, Tottori
University, and Nagoya University. Several universities carry
out research in grains processing.
The Korean Rural Development Administration of the Min-
istry of Agriculture, Food and Rural Affairs has a series of
national research institutes, including the National Institute
of Crops Science (www.nics.go.kr/english) with a focus on
rice, wheat, maize, soybean, and maize breeding, agronomy
and applied biotechnology, and postharvest technology;
National Academy of Agricultural Science in Suwon (www.
naas.go.kr/English/naas_main.asp) with relevant work on
crop protection and agro-ecology; National Institute of Agri-
cultural Biotechnology (www.niab.go.kr); and several provin-
cial agricultural and extension institutes. Grains research is
strongly rice-focussed, but with an increasing environmental
protection and biotechnology emphasis. Of the ten Flagship
Korean National Universities, eight have agriculture colleagues
that undertake grains research to varying extents, including
Chonbuk, Chonnam, Chungbuk, Chungnam, Gyeongsang,
Kangwon, Kyungpook, and Seoul National Universities.
China has the largest national agricultural research systems
(NARS) in the world and also the most geographically and
administratively decentralized. The government has placed a
high priority on developing research capacity and infrastruc-
ture over the last decade, and private sector expenditure in
research has also increased. Because of the large range of pro-
duction environments, research on both temperate and tropi-
cal grains is important, and some institutes undertake
pioneering grains research, for example, in the development
of high-yielding rice varieties, rice functional genomics, and
transgenic crops. In recent years, the Chinese government has
been making major investments in R&D physical infrastructure
and equipment, and there has been a move to retain or attract
back eminent scientists from the Chinese diaspora. There is an
increasing shift in emphasis in China from research aimed at
increasing grain production to increasing quality and ensuring
that production is environmentally sustainable, especially in
view of the increasing pressure on water resources for agricul-
ture in China.
Grains research is carried out in five main types of organi-
zations. The Chinese Academy of Agricultural Sciences (CAAS,
www.caas.net.cn/en) under the Ministry of Agriculture has the
largest number of grains research projects. There are 32
research institutes (employing over 5000 research staff directly
affiliated with CAAS; another nine are hosted jointly with
universities and provincial governments), with a growing
emphasis on environmental safety and sustainability, rather
than just higher yields. CAAS has recently consolidated its
vast array of research projects into eight disciplinary clusters.
The Crop Science crop germplasm resources cluster addresses
genetics and breeding, molecular biology, cultivation and
physiology, molecular breeding, functional genomics, proteo-
mics, metabonomics, and bioinformatics participating grains
institutes include the Institute of Crop Sciences, Biotechnology
Research Institute, China National Rice Research Institute, and
the Oil Crops Research Institute. The Agricultural Resources
and Environment cluster addresses plant diseases, insect pests,
weeds and rodents, dryland and water-efficient agriculture, soil
sciences, and plant nutrition, and participating grains institutes
include the Institute of Plant Protection, Institute of Environ-
ment and Sustainable Development in Agriculture, Biotech-
nology Research Institute, the Farmland Irrigation Research
Institute, China National Rice Research Institute, and the
Agro-Environmental Protection Institute. Grains research is
also undertaken within the Agricultural Mechanization and
Engineering; Agro-Product Quality, Safety, and Processing;
and Agricultural Information and Economics clusters. The
Agricultural Science and Technology Innovation Program
(ASTIP), launched in early 2013, funds four crosscutting objec-
tives supporting long-term and interdisciplinary research,
capacity building, expanding research support facilities and
infrastructure, and fostering international cooperation over
the next 13 years.
The Chinese Academy of Sciences (www.english.cas.cn),
under the Ministry of Science and Technology, also carries out
some research relating to grains, at the Institute of Botany and
Institute of Genetics and Developmental Biology (Beijing; basic
plant biology and biotechnology), Institute of Zoology (Beijing;
pest management), Institute for Soil Research (Nanjing),
National Centre for Genetic Research (Shanghai; rice genomics),
Chengdu Institute of Biology (cereal breeding), Institute of Soil
and Water Conservation (Yangling; agronomy), NW Institute of
Plateau Biology (Qinghai; crop assessment), and Shijiazhuang
Institute of Agricultural Modernization (Hebei; farming systems
for the North China plain) among others.
Several provincial academies of agricultural science have
strengths in grains research including Jiangsu, Sichuan, Guang-
dong, Fujian, Henan (for wheat), Shandong, Liaoning, and
Gansu. The State Administration of Grain (www.chinagrain.
gov.cn/english/Scientific%20Research.html) also carries out
grain storage research, including through the Grain Storage
Research Institute in Chengdu. In recent years, much of the
increase in research capacity has come from the universities.
Each province has at least one agricultural university, but the
three strongest universities in agricultural science are Nanjing
Agricultural University, Zhejiang University, and Huazhong
Agricultural University. Other agricultural universities carrying
out grains research are South China Agricultural University
(Guangzhou), Shenyang Agricultural University, Southwest
Agricultural University (Chongqing), and Northwest A&F Uni-
versity (Yangling). Further information on grain production
and consumption in China is discussed elsewhere in this
Encyclopedia.
South Asia
India also has one of the largest NARS in the world and a
traditional strength in crops research, particularly grain
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World
sciences, given the ongoing national focus on food security.
Most grains research is done within the Indian Council of
Agricultural Research (ICAR, www.icar.org.in) and state
agricultural universities. Of the 45 ICAR Central Research Insti-
tutes, most of the grains research institutes are within the Crop
Science Division. Major thrusts include development of grain
varieties resistant/tolerant to a range of abiotic and biotic
stresses, hybrid technology, application of molecular tech-
niques, plant genetic resources conservation, and farming sys-
tems research. The main ICAR institutes carrying out grains
research include
the Indian Agricultural Research Institute (emphasis on
more strategic research);
the Indian Institute of Pulses Research;
the National Bureau of Plant Genetic Resources;
the National Research Centres for Groundnut, Rapeseed
Mustard, Sorghum and Soybean;
the National Centre for Integrated Pest Management;
the National Research Centre on Plant Biotechnology;
the Central Institute of Post-Harvest Engineering and
Technology;
the Central Soil Salinity Research Institute;
the Central Arid Zone Research Institute;
the Central Research Institute for Dryland Agriculture;
the Indian Institute of Soil Science;
Directorates of Wheat Research, Sorghum Research, Oil-
seeds Research, Soybean Research, Rice Research, and
Weed Science Research.
Several smaller project directorates covering biological control,
oilseeds, rice, wheat, and maize. All India Coordinated
Research Projects attempt to link institutes as a mechanism to
build nationwide coordinated interdisciplinary research net-
works increasing the level of interinstitutional collaboration
is currently a high priority of the Indian government.
The second significant resource is the 55 state agricultural
universities (SAUs). Some of the SAUs with strong grains
research programs include Acharya NG Ranga AU (Andhra
Pradesh), CCS Haryana AU, GB Pant AU (Uttar Pradesh),
Punjab AU, and Tamil Nadu AU. The Council for Scientific
and Industrial Research (CSIR) has the Central Food Techno-
logical Research Institute (CFTRI, Mysore, www.cftri.com)
has a department that researches cereal (especially wheat)
milling, baking, and biochemistry. A number of private sector
companies undertake research into crop breeding and plant
protection. Further information on grain production and
consumption in India is discussed elsewhere in this
Encyclopedia.
Pakistan also has one of the largest numbers of grains
researchers in the developing world. The main institute of the
Pakistan Agricultural Research Council (PARC, www.parc.gov.
pk), the National Agricultural Research Centre in Islamabad,
covers soil fertility; wheat, rice, sorghum, and millet; farm
machinery; water resources; agricultural biotechnology; verte-
brate pests; crop diseases; crop production; land resources;
plant genetic resources; technology transfer; and seed sciences.
There are a separate tropical agricultural research center at
Karachi and four arid zone research centers in different regions
of Pakistan. There are several agricultural universities in
Pakistan that carry out grains research, the largest being the
447
Pakistan Agricultural University at Faisalabad. Sindh
Agriculture University, the University of Arid Agriculture
(Rawalpindi), and the Northwest Frontier Agricultural Univer-
sity (Peshawar) are all smaller, but with a similar range of
activities. The Ministry of Science and Technology manages
the Pakistan Council of Scientific and Industrial Research,
which carries out grains postharvest research. There is also a
separate National Institute for Biotechnology and Genetic Engi-
neering (Faisalabad) as well as the NARC Institute of Agricul-
tural Biotechnology and Genetic Resources. State departments
of agriculture in Balochistan, Punjab in particular, carry out
some grains research.
Elsewhere in South Asia, the Bangladesh Agricultural
Research Council has a central Bangladesh Agricultural
Research Institute (which carries out significant amounts of
grains research) together with satellite institutes covering soils
and rice. The largest university conducting grains research is
Bangladesh Agricultural University.
Southeast Asia
The major Indonesian government agricultural research and
development agency is the Indonesian Agency for Agricultural
Research and Development (IAARD, www.en.litbang.deptan.
go.id). Research institutes working on grains (unlike planta-
tion crops) are publicly funded and those focussing on grains
include the Centers for Food Crops R&D (Bogor, W Java), Rice
Research (Subang, W Java), Agricultural Engineering R&D
(Serpong, W Java), and Postharvest R&D (Bogor, W Java).
Typically smaller research institutes work on legume and
tuber crops (Malang, E Java), cereals (particularly maize)
(Makassar, S Sulawesi), and soils (Bogor, W Java). Adaptive
research is carried out by Assessment Institutes for Agricultural
Technology (under IAARD, but located in each province).
Major agricultural universities include Bogor Agricultural Uni-
versity (IPB), Gadjah Mada University, Hasanuddin University,
University of Mataram, Sam Ratulangi University, Sebelas
Maret University, Andalas University, Haluoleo University,
Brawijaya University, and Padjadjaran University.
Under the Philippine Department of Agriculture (DA),
there is a national rice research center (PhilRice in Munoz,
Luzon, www.philrice.gov.ph), addressing both research and
technology dissemination, and the Bureaus of Plant Industry,
Soil and Water Management (www.bswm.gov.ph), and Post-
harvest Research and Extension (which includes an emphasis
on grain storage research). The Food and Nutrition Research
Institute (www.fnri.dost.ph) and the research planning and
funding body, the Philippine Council for Agriculture, Aquatic
and Natural Resources Research and Development (www.
pcaarrd.dost.gov.ph), sit under the Department of Science
and Technology. There are also Regional Integrated Agricul-
tural Research Centers there is one organization in each of
the 15 regions, but some have 23 research stations and several
focus on grain crops. The universities have a far greater share of
research capacity than DA. The strongest university in agricul-
ture including grains research is the University of Philippines at
Los Banos, which has specialized institutes of plant breeding
and of farming systems research. These include Benguet
State University, Central Luzon State University, Visayas State
University College of Agriculture, Don Mariano Marcos State
448 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World
University, Central Mindanao University, and the University of
Southern Mindanao. There are also several others with some
strengths in agriculture (Mindanao State University, Bicol Uni-
versity, and Central Visayas State College). Some of the univer-
sities have regional research and development centers and also
have quite good extension/outreach systems. While significant
private sector research enterprises are present in horticulture,
grains research remains in the public domain.
The Thailand Department of Agriculture (DOA, www.doa.
go.th/en) is responsible for crops and farm mechanization
research, while the Department of Agricultural Extension
(DOAE) is responsible for transfer of technology to farmers.
While there are experiment stations in every region of the
country, most research is carried out in Bangkok. DOA has
programs covering agricultural chemistry, agricultural engi-
neering, plant pathology and microbiology, entomology and
zoology, soil science, botany, and weed science, crop protec-
tion, and specialist rice and field crops institutes (there is
reasonable research capacity in maize and legumes research).
The Land Development Department conducts research in rela-
tion to land improvement and soil and water conservation.
Grains postharvest technology is carried out within the
Thailand Institute of Scientific and Technological Research
(TISTR, www.tistr.or.th/tistr_en) in Bangkok. One of the
three research centers with National Science and Technology
Development Agency is Genetic Engineering and Biotechnol-
ogy (BIOTEC, www.biotec.or.th/en), which carries out some
rice biotechnology work. Kasetsart University in Bangkok,
Khon Kaen University, Chiang Mai University, Prince of
Songkla University (Hat Yai), and King Mongkuts Institute of
Technology in Bangkok are all involved in grains research, the
latter having a special focus on grains postharvest technology.
In Malaysia, agricultural R&D spending as a percentage of
production value (research intensity) remains comparatively
high. The Rice and Industrial Crops Research Centre of the
Malaysian Agricultural Research and Development Institute
(MARDI, www.mardi.gov.my) researches rice, maize, and
groundnuts as well as several nongrain crops. The most rele-
vant activity is development of high-quality rice varieties with
disease resistance and accompanying agronomy. The food
technology research center does work on the development of
rice-based snacks and other food products. The major univer-
sities carrying out grains science research is the Universiti Putra
Malaysia and Universiti Kebangsaan Malaysia.
Most grains research in Vietnam is conducted under
the Ministry of Agriculture and Rural Development (MARD,
www.agroviet.gov.vn). Other relevant ministries are education
and training (university sector), science and technology (www.
most.gov.vn), which has some excellent crop biotechnology
and plant protection research facilities, and the Ministry of
Industry (which has an Oil Plant Research Institute with
the mandate for peanut, soybean, sunflower, and sesame).
While most institutes are located in either Hanoi or Ho Chi
Minh City, many maintain research centers in more remote
locations. The key MARD institutes in grain science are the
Food Crops Research Institute, National Institute for Plant
Protection, National Institute for Soils and Fertilizers, Institute
of Agricultural Engineering and Post Harvest Technology, Insti-
tute of Agricultural Science in the South, Vietnam Agricultural
Science Institute, Agricultural Genetics Institute, National
Maize Research Institute, and Mekong Rice Research Institute.
The Department of Plant Protection of MARD is also becoming
an important applied research organization. The main univer-
sities with research interests in grain science include Hanoi
Agricultural University, Hanoi University of Science (also
known as Hanoi College of Sciences), Nong Lam University,
Hue University of Agriculture and Forestry, Thai Nguyen Uni-
versity, and Can Tho University. Further information on grain
production and consumption in Southeast Asia is discussed
elsewhere in this Encyclopedia.
Central and South America
The major grains R&D capacity in Central America is within
Mexico. Initially, it was focussed in several of the Instituto
Nacional de Investigaciones Forestales, Agricolas y Pecuarias
(INIFAP, www.inifap.gob.mx) crops institutes based on differ-
ent agroecological zones (www.inifap.gob.mx), but it is now
spread across a range of universities and other government
agencies. Grains food research and biotechnology is carried
out in the Consejo Nacional de Ciencia y Tecnologia
(CONACYT, www.ciad.mx). Grains research is carried out in
the Universidad Autonoma Chapingo, the Instituto Polit-
ecnico Nacional, and over a dozen regional universities.
In South America, Brazil has a well-developed NARS, with
more agricultural researchers than any other country in Latin
America. Many centers fall under EMBRAPA (Empresa Brasile-
ira de Pesquisa Agropecuaria, www.embrapa.br), a specialized
science agency of the Agriculture Ministry. Brazil has the largest
NARS of any Central or South American country, followed by
Mexico; Brazil also has an advanced university system. In addi-
tion, 16 of Brazils 26 states operate agricultural research insti-
tutes, with Sao Paulo the largest. There are separate EMBRAPA
institutes relating to genetic resources and biotechnology
(CENARGEN), upland rice and beans (CNPAF), corn and sor-
ghum biotechnology (CNPMS), soils technology transfer
(CNPS), soybean and sunflower (CNPSO), wheat, barley,
oats, soybeans, and biotechnology (CNPT), temperate climate
agricultural research (CPACT, irrigated rice), and food technol-
ogy (CTAA), as well as at least 12 regional centers. There are
also nonprofit centers Foundation Center for Wheat Experi-
mentation and Research (FUNDACEP, www.fundacep.com.
br), the Central Agricultural Cooperative of Technology Devel-
opment and Economics (COODETEC; soy, maize, and wheat),
and the Rice Institute of the Rio Grande do Sul (IRGA). Several
universities carry out grains research the federal universities
of Goias (www.agro.ufg.br), Lavras, Paraiba, Pelotas (www.
faem.upfel.edu.br), Rio Grande do Sul, Rural do Rio de Janeiro
(www.la.ufrrj.br), Santa Catarina (www.cav.udesc.br), Sao
Carlos (www2.ufscar.br), Santa Maria, Vicosa, and Brasilia as
well as several state universities (Santa Catarina State, Paulista,
Campinas, Sul de Santa, and Sao Paulo). Some Brazilian com-
panies, such as Agroceres, DuraFlora, Mogiana, and Braskalb,
carry out breeding and agronomy research to support seed
distribution businesses.
The Argentine Government Instituto Nacional de Tec-
nologia Agropecuaria (INTA, www.inta.gov.ar) is under the
Agriculture Ministry, and capacity has increased significantly
over the last decade due to significant increase in government
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World
investment. There are also a large number of programs and
centers relevant to grains research under CONICET (Consejo
Nacional de Investigaciones Cientificas y Tecnicas, www.con-
icet.com.ar), which is under the Culture and Education Minis-
try. This includes a number of crop breeding, agronomy, and
natural resource (CADIC and CEFYBO), plant protection
(CEAVE and IFEVA), postharvest technology (CEFOBI and
CRILAR), and plant physiology and molecular biology
(INFIVE and INGEBI) centers. Several national universities
have agronomy science faculties Buenos Aires, Buenos Aires
Province, Catamarca, Comahue, Cordoba, Cuyo, Entre Rios,
Jujuy, La Pampa, La Plata, Litoral, Lomas de Zamora, Mar del
Plata, Noredeste, Rio Cuarto, Rosario, Santiago del Estero, de
Sur, de Salta, Tucuman, and de Formosa.
The Chilean Ministry of Agriculture has an Instituto de
Investigaciones Agropecuarias (INIA, www.inia.cl), which
undertakes some grains research, although overall agricultural
research is much more limited than in Brazil or Argentina. There
are also agronomy departments at several universities Austral
University (www.agrarias.uach.cl), the University of Conception
(www.chillan.udec.cl), the University of La Frontera, the
University of La Serena (www.userena.cl), the University of
Magallanes, the University of Talca, the University of Tarapaca
(www.uta.cl), Universidad de Chile, and Universidad Ibero-
americana de Ciencias as well as the Catholic universities
(Temuco (www.uctemuco.cl), Valparaiso (www.ucv.cl), and
Maule). In Colombia, the main government crops research
agency is CORPOICA (www.corpoica.gov.co); but much
more research is now carried out through the producer associa-
tion, Asociacion Nacional de Acuicultores de Colombia
(ACUANAL); there are research groups working on rice. Apart
from the campuses of the national university in Bogota, other
universities with grains agronomy faculties include Cordoba
(www.unicordoba.edu.co) and Tolima (www.ut.edu.co). The
main research agency in the Ecuador agriculture ministry is
INIAP (Instituto Nacional Autonomo de Investigaciones Agro-
pecuarias); universities in most regional centers have small
agronomy departments. In Uruguay, the government Instituto
Nacional de Investigacion Agropecuaria (INIA, www.inia.org.
uy) is the largest crops research institute. Further information
on grain production and consumption in South America is
discussed elsewhere in this Encyclopedia.
Africa and the Middle East
South Africa has the best-developed NARS of the African
countries, although its capacity has declined over the last
decade. South Africa maintains reasonable research capacity
in maize (the main grain crop), sorghum, and some experi-
ence in wheat and a little in barley. The Agricultural Research
Council (www.arc.agric.za) is the main agricultural research
structure and has a series of research institutes, several carry-
ing out grains research particularly the Grain Crops Institute
and the Small Grains Institute, with some other institutes
carrying out supporting natural resource management
research. The nine provincial departments also carry out
selection and agronomy research to varying degrees. There
are some well-resourced grains research programs at the Uni-
versities of Pretoria (www.web.ip.ac.za), KwaZulu-Natal
449
(www.sasa.ukzn.ac.za), the Free State (www.ufs.ac.za), and
Stellenbosch (www.acadmic.sun.ac.za/agric/). In the 1990s,
there has been a significant shift in research emphasis in
South Africa to accommodate the needs of the wider popula-
tion, rather than just white commercial agriculture, to include
emerging farmers and subsistence farmers in former home-
lands. Private companies such as Monsanto and Pannar carry
out seed and fertilizer research.
In Zimbabwe, most crops research is carried out by the
Department of Research and Specialist Services in the Minis-
try of Lands (www.drss.gov.zw) and Agriculture, Mechani-
zation and Irrigation Development, with the University of
Zimbabwe (www.uz,az.zw) maintaining reasonable strength.
The Kenya Agricultural Research Institute (KARI, www.kari.
org) is comparatively strong in crops and livestock and the
Kenyan government agricultural research system one of the
strongest in Africa. In the grains area, mandate crops include
cereals (wheat, maize, sorghum, millets, and rice) and grain
legumes with efforts ranging from plant breeding for adapta-
tion to different agroecological zones and stress resistances,
agronomy, postharvest technology, and cooking quality.
Almost all Kenyan grain production is rain-fed. The breeding
and postharvest research is backed up by a soil and water
management research program. The Kenya Industrial R&D
Institute (www.kirdi.go.ke) also carries out some postharvest
research. Some grains research is carried out at various uni-
versities (Jomo Kenyatta University of Agriculture and Tech-
nology (www.jkuat.ac.ke), Egerton University (www.egerton.
ac.ke), Moi University (www.mu.ac.ke), the University of
Eastern Africa (www.uaeb.ac.ke), and the University of Nai-
robi (www.ounbi.ac.ke). The Ethiopian Institute of
Agricultural Research (www.eiar.gov.et) has national and
regional centers emphasizing maize, plant protection, soils,
and agricultural mechanization. Crops research is carried
out at Hawassa University (www.hu.edu.et), Haramaya
University (www.haramana.edu.et), Jimma University (www.
ju.edu.et), and Mekelle University (www.mu.edu.et). In
Uganda, most grains research capacity is within the main
government National Agricultural Research Organization
(NARO, www.naro.go.ug) and at Makerere University (www.
agric.mak.ac.ug).
In Nigeria, the Ministry of Agriculture through the Agricul-
tural Research Council of Nigeria (www.arcnigeria.org) has a
series of research institutes National Cereals Research Insti-
tute, Lake Chad Research Institute (www.icresmaid.org), and
the Institute for Agricultural Research (www.abu.edu.ng). Post-
harvest technology research is carried out under the Ministry of
Industry at the National Stored Products Research Institute
(www.nspiilofin.com), while the Ministry of Science and Tech-
nology has established a National Centre for Genetic Resources
and Biotechnology (www.nacgrab.gov.ng). Universities with
relevant research programs include the University of Ibadan
(www.agric.ui.edu.ng), University of Nigeria (www.unn.edu.
ng), Bayero University, Ahmadu Bello University (www.abu.
edu.ng), and Obafemi Awolowo University (www.oauife.edu.
ng). In Zambia, the Zambia Agricultural Research Institute
(www.zari.gov.zm), Postharvest Food Processing and Nutri-
tion Institute (www.nsirir.org.zm), and the University of Zam-
bia (www.unza.zm) are the main grains research providers. In
Tanzania, it is the Directorate of Research and Development of
450 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Research Organisations of the World
the Ministry of Agriculture and Food Security (www.agricul-
ture.go.tz) along with the Sokoine University of Agriculture
(www.suanet.az.tz); in Botswana, the Department of Agricul-
tural Research (www.dar.gov.bw), while in Ghana, the Crops
Research Institute of the Council for Scientific and Industrial
Research (www.cropsresearch.org), along with the University
of Ghana (www.ug.edu.gh) and the Kwame Nkrumah Univer-
sity of Science and Technology (www.knust.edu.gh), are the
main crops research providers. Further information on grain
production and consumption in Africa is discussed elsewhere
in this Encyclopedia.
The Israeli NARS is reasonably well resourced. Within the
Agricultural Research Organization (www.agri.gov.il), the
Institute for Plant Sciences has programs on cereal crop agron-
omy to overcome drought, disease, and heat stress. There are
separate institutes for plant sciences, soil water and
environmental sciences, postharvest and food sciences, and
agricultural engineering, which also carry out grains-related
research. Universities carrying out grains research include
Technicon (Israel Institute of Technology (Haifa)) and the
Ben-Gurion University of Negev.
Egypt also has a large government agricultural research
system, within the Agricultural Research Center (ARC, www.
arc.sci.eg), within the Ministry of Agriculture and Land Recla-
mation. In ARC, there are separate programs for plant protec-
tion, plant pathology, food technology (bread and pasta
research), agricultural genetic engineering, and farming sys-
tems research relating to wheat, barley, maize, rice, sorghum,
legumes, and oil crop production. Universities with agriculture
faculties include Alexandria, Cairo, Ain Shams, Al-Azhar, Suez
Canal, Zagazig, Assiut, Mansoura, Minia, Menoufia, Tanta, and
South Valley. Further information on grains research in other
West Asian and North African countries is discussed elsewhere
in this Encyclopedia.
Exercises for Revision
What is the largest national agricultural research system in
the world?
In what regions of the world is grains research capacity
thought to have decreased over the last decade?
Name three countries where the involvement of the private
sector in grains breeding has increased significantly over the
last decade.
What role does the national Australian government have in
grains R&D delivery?
What are the two major organizations or systems undertak-
ing grains research in India?
Exercises for Readers to Explore the Topic Further
How do agricultural research in Brazil contributed to the
export success of its grains industries?
How do the priorities of grains research for the Indian state
agricultural universities differ when they are based in dif-
ferent geographic zones of India?
Being a small country, why doesnt New Zealand simply
adopt the grain varieties and agronomy research findings of
its larger neighbor, Australia?
See also: Agronomy of Grain Growing: Implication of Climate
Changes; Wheat: Agronomy; Breeding of Grains: Barley: Genetics
and Breeding; Canola/Rapeseed: Genetics and Breeding; Maize:
Breeding; Rice: Breeding; Soybean: Germplasm, Breeding, and
Genetics; Wheat Breeding: Exploiting and Fixing Genetic Variation by
Selection and Evaluation; Food Grains and the Consumer: Cultural
Differences in Processing and Consumption; Grains Around the
World: Grain Production and Consumption: China and South-East
Asia; Grain Production and Consumption in Oceania: Australia and
Pacific Countries; Grain Production and Consumption: Overview; Grain
Production and Consumption: South America; Grain Production and
Consumption: Africa; Production and Consumption of Grains: India;
Scientific Organizations Related to Grains: Scientific Societies
Associated with Grain Science; The Basics: The Grain Crops: An
Overview; The Cereal Grains: An Overview of the Family of Cereal
Grains Prominent in World Agriculture; Wheat-Based Foods:
Cereals: Overview of Uses: Accent on Wheat Grain.
Further Reading
Beintema NM and Stads G-J (2010) Public agricultural R&D investments and capacities
in developing countries: Recent evidence for 2000 and beyond. Background note for
the Global Conference on Agricultural Research for Development, Montpellier,
France, March 2831.
Beintema N and Stads G-J (2011) African Agricultural R&D in the New Millennium:
Progress for Some, Challenges for Many. Washington, DC: International Food
Policy Research Institute.
Cassas J, Sohl M, and Hafaz H (1999) The National Agricultural Research Systems in
the West Asia and North Africa Region. Aleppo, Syria: ICARDA, p. 278.
Corke H and Cai YZ (2016) Grain production and consumption: China and South East
Asia. In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of
Food Grains, 2nd edn., vol. 1, pp. 374382. Elsevier Ltd.
Flaherty K, Stads G-J, and Srinivasacharyulu A (2013) Benchmarking Agricultural
Research Indicators Across Asia-Pacific. Washington, DC: International Food Policy
Research Institute.
Khatkar BS, Chaudhary N, and Dangi P (2016) Production and Consumption of Grains:
India. In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of
Food Grains, 2nd edn., vol. 1, pp. 367373. Elsevier Ltd.
Pardey PG and Alston JM (2006) Agricultural R&D in the Developing World: Too Little,
Too Late? Washington, DC: International Food Policy Research Institute.
Roseboom J, Cremers M, and Lauckner B (2001) Agricultural R&D in the Caribbean: An
Institutional and Statistical Profile. The Hague: International Service for National
Agricultural Research, ISNAR Research Report No. 19.
Taylor JRN (2016) Grain production and consumption, Africa. In: Corke H, Wrigley C,
Faubion J, and Seetharaman K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1,
pp. 359366. Elsevier Ltd.
Wrigley CW and French RJ (2016) Grain production and consumption in Oceania:
Australia and Pacific Countries. In: Corke H, Wrigley C, Faubion J, and Seetharaman
K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1, pp. 409419. Elsevier Ltd.
 Research Organizations of the World: Global Trends and the Commercial Sector
JH Skerritt, University of Queensland, St. Lucia, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Greater emphasis on demand-driven rather than science-
driven research
Increasing emphasis on topics such as production sustain-
ability, croplivestock systems, systems modeling, and the
use of techniques from molecular and information sciences
but also a return to treatment of research on grain produc-
tivity and yield.
Specialization into particular areas common as well as
clearer identification by funders and institutes about how
near to market
Greater collaboration between disciplines and between
research institutes as well as increased private sector has
greater involvement in agricultural R&D
Major commercial groups involved in grains and grain
product research
Learning Objectives
To obtain a better understanding of global trends in grains
research and grains research undertaken by the commercial
(biotechnology, seed, crop protection, milling and baking,
brewing, and cereal foods) sector.
Grains Research in the Commercial Sector
The private sector has greater involvement in agricultural R&D,
especially in linkages between seed business and biotechnol-
ogy/life science concerns. Pardey and Beintema had calculated
that since the mid-1990s, in developed countries, expenditures
from the private sector in agricultural research have exceeded
public expenditure.
While bakery and grains processing companies have always
engaged in research to support new product development, a
particular focus of their research over the last decade has been
in the development of grain food products that promote health
and well-being. Despite the health benefits, there are chal-
lenges to increasing consumption of whole grain foods includ-
ing taste preferences and convenience of these foods. In
addition, there is a growing tendency for increasing numbers
of people to avoid cereals in their diets because of perceived
health issues, even though the documented prevalence of
celiac disease and cereal allergy is comparatively low. Develop-
ment and marketing of gluten-free foods has thus become a
significant business opportunity.
Grains Processing Companies
Most of the research carried out by these companies is con-
cerned with practical applications (e.g., new product
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00058-9
development) aimed to support their flour milling and bakery
product businesses (including breakfast cereals), but some
carry out more basic research and their staff contribute actively
to research conferences and the scientific literature. Major
examples in the United States include ConAgra Foods in
Omaha, NE (www.conagrafoods.com), General Mills in Min-
neapolis, MN (www.generalmills.com), Kellogg Company in
Battle Creek, MI (www.kelloggcompany.com), and Nabisco,
New Jersey (www.snackworks.com). Nabisco Research carries
out fundamental studies on dough rheology and physical
properties. Riceland Foods (www.riceland.com, Stuttgart, AR,
the United States) and Riviana Foods (www.riviana.com,
Houston, TX, the United States) are two large manufacturers
of rice- and soy-based food ingredients and undertake applied
research to support their markets. The National Starch and
Chemical Company, Bridgewater, NJ (www.nationalstarch.
com), focuses on cereal starch-based polymers and supports a
significant research program including work on new adhesives,
specialty food starches, and biodegradable packaging mate-
rials. Cargill has wheat flour milling and maize wet milling
divisions and provides grains-based feeds for livestock.
Through Cargill Health and Food Technology (www.cargill-
foods.com), it focuses on nutraceuticals research, especially
on developing products derived from soy.
In Japan, two large milling companies, Nisshin Flour Mill-
ing (www.nisshin.com/english/group) and Nippon Flour
(www.nippn.com), have played an important role over the
last few decades in increasing the consumption of wheat in
Japan. They also carry out a significant amount of research. In
the United Kingdom, Rank Hovis (www.rankhovis.co.uk),
ADM Milling (www.adm.com/en-US/MillingUK/Pages/
default.aspx), Associated British Foods, Allied Bakeries, and
British Bakeries are among the largest milling and baking
companies, and they conduct some in-house research and
technology development. Barilla (www.barillagroup.com/cor-
porate/en/home/lavora-con-noi/organization-and-job-fami-
lies/research-development-quality.html) has a large cereal
products research center in Italy, while in Australia, Goodman
Fielder (www.goodmanfielder.com.au) and GWF (www.geor-
gewestonfoods.com.au) have active technology development
programs. Other companies within the oilseed crushing and
animal feeds industry also have applied research programs.
Baking Industry Associations
Industry associations are also an important provider of
research and technical services in the grains industry. In the
United Kingdom, Campden BRI (www.campdenbri.co.uk),
Chipping Campden (Gloucestershire) and Nuffield (Surrey),
is a membership-based organization that provides research
and consultancy services for cereal-processing companies.
Leatherhead Food Research (www.leatherheadfood.com)
started as a UK food-industry association but has evolved
451
452 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Global Trends and the Commercial Sector
into an independent provider of food research and market
information. In France, ARVALIS Institut du vegetal, a farmers
organization, carries out research on grains to satisfy the
demands of markets and consumers. AIB International
(www.aibonline.org), in Manhattan, KS, the United States,
was founded by the North American baking industry to apply
research findings to the industry in the United States and
internationally. Emphasis of the research includes cereal sci-
ence, baking technology, nutrition, and food safety.
BRI Australia Ltd. (formerly the Bread Research Institute)
has now become part of GrainGrowers (www.graingrowers.
com.au), an independent Australian organization supported
by the industry through membership fees and by competitive
government and industry research grants. It carries out applied
research in flour milling, baking, and Asian foods. The Japan
Institute of Baking Technology in Tokyo also plays a similar
role.
Seed and Grain Biotechnology Companies
In a large number of cases, these two sectors have merged since
the 1990s. A brief description of some of the largest multina-
tional companies is in Table 1. Many of the companies were
initially agrochemical suppliers, but the development of trans-
genic, herbicide-resistant grain crops provided them with an
opportunity to market the seed and agrochemical together as a
package. Several companies have an obvious leadership role in
the development of agrochemicals and transgenic crops. How-
ever, the merger of many of these enterprises with seed com-
panies increased their importance in breeding and agronomy
and, more significantly, resulted in the consumer benefiting
from research on transgenic crops. Genetic engineering has
been carried out mostly on maize, cotton, soybean, and canola
with limited commercial progress with barley and wheat
because of consumer resistance and in some cases technical
challenges. To achieve resistance to herbicides and insecticidal
activity, genes for Bacillus thuringiensis toxin were incorporated.
For example, Monsanto Companys products (St. Louis, MO)
include herbicide-resistant soybeans, canola, and corn and
insect-protected corn. Several of the companies also have
broader genomics and gene-discovery programs. Refer to the
papers by King and OBrien for reviews of barley and wheat
breeding, respectively, and by Gill and Henry for reviews of
genome mapping and genomics, respectively.
Research and development budgets of several of these com-
panies run into several hundreds of millions of dollars, com-
parable with the government agricultural R&D budgets in
some medium-sized developed countries. Over the last decade,
early commercial impact in the cotton industry has translated
into significant market share for grains, particularly maize and
soybean. Consumer resistance and technical challenges mean
that significant commercialization of genetically modified
wheat and barley is lagging. Industry describes the products
of biotechnology as seed with trait meaning traits such as
herbicide or insecticide resistance. Of these, herbicide resis-
tance (enabling particular herbicides to be sprayed in a stand-
ing grain crop) is the main trait for relevant grain crops, while
insect resistance through B. thuringiensis (Bt) technology is
second in importance. Some other stacked traits provide tol-
erance to abiotic stresses, diseases, and control pollination.
Brewing Companies
These form a second large group of private sector companies
carrying out grains research. The largest brewing industry sup-
ported research organization is Campden BRI (www.campden-
bri.co.uk) in the United Kingdom, which was formed out of
merger between the Campden and Chorleywood Food Research
Association and Brewing Research International. It undertakes
research on behalf of members worldwide on aspects of
food safety, barley and malt quality, fermentation, and beer
quality. In addition, a number of the major brewery companies
have significant in-house R&D activities, such as the giant
USEuropean brewing conglomerate ABInBev, SABMiller in
the United Kingdom, and Tepral (Strasbourg, France) as well
as smaller brewing companies worldwide. Applied research on
barley and malt composition and quality is carried out.
The research activities of Carlsberg Brewers spread much
further than malting and brewing. Since the establishment of
research laboratories in 1875, Carlsberg has made major contri-
butions to fundamental chemistry and such as the development
of the Kjeldahl protein analytic method. The Carlsberg Labora-
tory (www.carlsberglab.dk) emphasizes both basic research and
applied research on barley grain protein and carbohydrates and
their reactions during brewing and genetic and biochemical
studies on amino acid transport, protein-folding seed germina-
tion, barley mutants, starch synthesis, and yeast physiology.
The Private Sector in Asia and Africa
Over the last decade, there has been increasing recognition by
governments and by aid donors that private sector involve-
ment in research can impact on poverty and increase produc-
tion in Asia and Africa. While poorly developed compared with
the United States, Europe, and AustraliaNZ, despite chal-
lenges with regulatory and policy settings, private company
involvement in applied grains research in varietal development
and selection, crop protection, and crop processing has signif-
icantly increased in recent years.
Indian private sector agribusiness investment is now quite
significant, being over $250 million in 2008/09. This includes
biotechnology and seed systems R&D to develop and dissem-
inate maize, wheat and rice cultivars, and rice, pearl millet, and
sorghum hybrids that provide superior yield under challenging
conditions. Private sector investment in African applied grains
R&D has also increased over the last decade and is growing
rapidly, coinciding with the development of more favorable
policy environments at a time when public funding for
research has stagnated, in several sub-Saharan countries. The
primary focus is the seed industry and plant variety develop-
ment, in particular maize (both proprietary and genetically
modified hybrids). In addition, private sector-bred varieties
of wheat, sorghum, barley, soybeans, and sunflower have dom-
inated over publicly bred varieties of the same species in terms
of recent crop registrations for the Republic of South Africa.
Major Trends among Grains Research Organizations
Improvements in grain crop yields over the last five decades are
the overwhelming reasons that despite a 120% increase in
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Global Trends and the Commercial Sector 453

Table 1 Major international life sciences and seed companies carrying out grains research

Company/locations Product/technology R&D emphasis Website

Monsanto Company Maize, soybean breeding, hybrids, herbicide, insect and wilt resistance, agrochemicals, seeds, www.monsanto.com
St. Louis, MO, USA and genomics
Agracetus (part of Crop plant transformation, therapeutic protein expression in plants www.agracetus.com
Monsanto)
Middleton, WI, USA
ADM (Archer Daniels Food ingredients; oilseeds; soy-based foods; feed ingredients; bakery science; nutraceuticals; www.adm.com
Midland) biofuels; extrusion research
Decatur, IL
Pioneer Hi-Bred Seeds, grain additives www.pioneer.com
(a Du Pont Company) Crop management, precision farming
Des Moines, IA, USA Grain seeds, especially hybrid maize (also millet, rice, sorghum, wheat)
BASF Crop protection products www.basf.de/en;
Limburgerhof, Germany Transgenic crops with stress tolerance, high oil content, and pathogen tolerance www.basf.com
Raleigh, NC, USA
Bayer Crop Science Crop protection www.
Akeno and Yuki, Japan bayercropscience.
Frankfurt, Germany com
Kansas City, MO, USA
Lyon, France
Syngenta Crop protection, seeds, and crop genetics www.syngenta.com
Jealotts Hill, UK Genomics, biotic stress tolerance, fungal diseases, insect and nematode control, marker-
Research Triangle Park, assisted breeding, maize, rice and soybean breeding, and transgenics
USA
Saint-Sauveur, France
Uberlandia, Brazil
Beijing, China
Dow Agrosciences Plant genetics and biotechnology; agrochemicals; seeds (hybrid and GM), traits (sunflower, www.dowagro.com
Indianapolis, IN, USA corn, and soybean) and oils; crop protection
Mycogen Breeding of sorghum, hybrid corn, sunflower, soybean, and canola www.mycogen.com
Indianapolis, IN, USA
Cargill Cereals for animal feed, biofuels; novel traits; oilseed breeding; biotechnology www.cargill.com
Ft Collins, CO, USA
Elk River MN, USA
Minneapolis, MN, USA
Vilvoorde, Belgium
KWS UK Wheat, barley, maize, and canola varieties www.kws-uk.com
Thriplow, UK
Limagrain Cereal breeding; seeds; plant genome analysis; transgenesis; biotechnology; maize, wheat, www.limagrain.com
Auvergne, France rapeseed, sunflower, soybean, barley, peas; baking technology
Biogemma Maize, wheat, rapeseed, sunflower breeding plant genome analysis; drought tolerance; www.biogemma.fr
Clermont Ferrand and disease resistance
Mondonville, France

global populations, food security is higher than ever. This is

because grain yields almost tripled in the same time, this
growing faster than population (http://faostat.fao.org). Suc-
cessful grains research, together with irrigation in favored
areas and favorable policy setting, is largely responsible for
averting starvation over the last half century.
As emphasized in the articles on taxonomy of grains
research systems, the last few years have seen a reemphasis of
food security, particularly for developing countries, and thus a
return to treatment of research on grain productivity and yield
as a priority. Also, specialization into particular areas some-
times driven in a top-down sense by funding bodies and
clearer identification by funders and institutes about how near
to market they are, that is, whether the institutes focus is on
variety development, precompetitive applied research, or
enabling plant science, have emerged as trends.
Increased Planning and Collaboration
Since the 1980s, there have been significant changes in the
organization and management of grains research. There has
been a greater degree of planning and an increased degree of
focus in the programs of many research organizations. Much of
the increased focus has been on market needs it is hard to
criticize the important influence this has had on breeding pro-
grams, grains chemistry, and processing research. Greater link-
ages between research on grain production and marketing are
important so that the research facilitates farmers who engage in
market-specified production, as opposed to research to under-
pin increased production of grain as a bulk commodity.
Over the last couple of decades, research centers have often
moved from being provided with guaranteed government allo-
cations to being dependent on competitive funding. Research
454 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Global Trends and the Commercial Sector

planning exercises have been in favor in a number of commer- Changes in Research Subject Matter
cial and government organizations across the globe. It is unclear
Grains agronomy programs have had to develop a stronger
to what extent individual creativity has been affected by these
research emphasis on sustainability of production in recent
trends certainly, there may be less scope for curiosity-driven
years, responding to a shift in community values as well as serious
grains research. The greater focus on markets by farmers and
scientific concerns about the long-term impact of some grain
producing for a particular market is reflected in research.
farming practices. Concerns about effects of irrigation, salinity,
An overarching trend has been the greater level of cooper-
acidification (especially with certain legume crops), loss of soil,
ation between research organizations; funding networks such
and of soil structure have led to an increase in research on reduced
as those established by the European Commission and Austra-
tillage, water conservation, and weed control and maintenance of
lian Government CRC system have facilitated this trend, as
soil fertility. Molecular breeding approaches have been imple-
well as organizational changes in major government depart-
mented through the development and release of new varieties a
ments such as US Department of Agriculture. However, it is
move from the laboratory to the field over the last decade.
probably true that US grains research is less well coordinated at
Impacts on climate variability and change and adaptation
a national level although locally, the close interactions
to variable climates have proved a significant focus for grains
between a number of USDA-ARS facilities and land grant uni-
research over the last decade. However, the shift in research
versities are very productive. The increased importance of net-
resources from productivity to other areas has been controver-
works in many areas of science is a response to the increased
sial Alston et al. demonstrated that a shift in research invest-
cost of carrying out research plus an increasing realization that
ment in productivity or yield-related outcomes has coincided
problems are multidimensional and are best addressed
with a plateauing in global yields for major cereals. Indeed, the
through multidisciplinary approaches. There has also been
major factor underpinning sustainable agriculture has been the
pressure from governments and funding bodies for research
ability to feed the worlds increasing population through
providers to network more, in part to avoid duplication of
enhanced yields, and with the exception of a few communities,
research and also to ensure that particular subprojects are
grains, especially cereals, are the central element of almost
carried out by the most qualified group. In the developing
diets. Since the 1940s, there has not been a significant increase
world, development banks such as the World Bank and Asian
in the area of land available for agriculture. However, it is true
Development Bank have funded reform programs that encour-
that with the expansion of irrigation systems and with new
age rationalization in the number of research providers as well
varieties and increased fertilizer use in the middle of the twen-
as greater coordination. In addition, since the spike in global
tieth century, there was an expansion of grain-growing areas.
grains prices in 200708, development agencies and banks
Since the 1980s, the expansion of irrigated areas has largely
have had a resurgence of interest in funding agriculture includ-
ceased and so production increases must come from increases
ing grains-related research.
in yield. In suitable ecologies, there has been a shift toward
Scientist-to-scientist collaboration across institutional and
more frequent harvests, especially in irrigated areas that may
national boundaries has been greatly facilitated by the commu-
accommodate three irrigated rice crops per year, as well as also
nications revolution the ability to e-mail (or more recently, use
improved rainfed rotations, for example, between a winter
cloud computing for) large amounts of data and draft manu-
cereal and a legume, rather than leaving the soil fallow over
scripts for immediate receipt and prompt publication online
summer. On the other with the recognition that breeding
has totally changed the possibilities for collaboration compared
targeted toward ideal high yield environments does not pro-
with 20 years ago. Globalization in industry through both
vide varieties that perform well under normal farm conditions,
development of alliances between commercial research pro-
a greater focus on breeding for representative environments
viders in different countries and increased emphasis on grains
and on abiotic stress tolerance has emerged.
breeding for export markets has also become a more impor-
Therefore, the key contributor to sustainability in grain
tant force. Markets are becoming more differentiated so there
production will continue to be productivity; increases are
is an increased emphasis on grain quality with targets for
important for both developing and developed country farmers,
breeding efforts being additional yield and disease resistance
as the terms of trade for farmers continue to become poorer. An
and on research aimed at improving the understanding of the
even greater challenge to developing countries is how North
genetic, chemical, and structural factors underpinning grain
American and European governments shield their grain
quality.
farmers from market realities through subsidies.
An increased focus on impact assessment, better under-
Livestock are becoming a very significant consumer of
standing of the concept of spillover but recognition of the lag
grains worldwide; the trend is especially noticeable in devel-
between successful R&D and adoption claimed to be 1525
oping countries, where meat consumption has increased sig-
years for much public agricultural R&D. While such lag periods
nificantly through the 1980s and 1990s, from a low base.
have been documented for early and mid-twentieth-century
Many of these animals are now grain-fed, so research has had
innovations such as hybrid maize and semidwarf cereals,
to underpin the increased importance of grains for feed rather
respectively, the lag period may persist it is two decades
than food applications. This research is also driven by
since the first genetically engineered grain crops were grown
increased intensification of monogastric animal production,
commercially, and yet for food grains, these still represent only
including in developing countries. In many countries, farmers
a minority of grains produced globally. So grains research
are not solely grain growers or cattle or sheep farmers, but
requires a longer-term view to be taken, both by aid donors
manage a mixed enterprise. There has been a more active
who fund grains R&D in developing countries through
development of systems-based research to address their needs.
commercial investors in advanced economies.
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Global Trends and the Commercial Sector
Another grains research area focused on Africa and South
and South East Asia is biofortification. There are several inter-
national projects focused on biofortification of staple cereals
with micronutrients such provitamin A, iron, and zinc. Some
approaches involve the use of genetic engineering technology
to transfer biosynthetic genes (e.g., for vitamins) into the grain
species, while other programs focus on exploration of natural
variation for the desired micronutrient in a grain species or in
primitive relatives of a target grain species followed by intro-
gression of the desired characteristics into high-yielding, high-
quality locally adapted varieties.
Consumer trends provide critical direction for many of the
commercial milling and baking organizations that conduct
research, including on product development. The trend toward
cooking less at home and convenience foods is continuing;
there is increasing popularity of gluten-free diets (whether or
not they are indicated on medical grounds) and greater con-
sumption of weight loss and diet snacks that have improved
taste. In addition, a focus on high-protein/low-fat and low-
carbohydrate grain products and emergence or niches for some
ancient grains such as spelt and nongrains/pseudocereals such
as quinoa and buckwheat used in traditionally grain foods
have also driven industry cereal foods product research and
development. In parallel, whole grain and whole fiber, spe-
cialty baked goods, have emerged as an increasingly important
sector for bakery research. Biofuel research continues to be
important in certain countries as demands grow for example,
in 2010/11, 40% of the US corn crop and 14% of soybean oil
were used for biofuels in the United States.
Institutional Trends
Another major change in the nature of agricultural research
since the 1980s, particularly in the United States and Europe,
has been the shift in balance of research to a greatly increased
role for the private sector. And within government systems, the
greatly increased resourcing and impact of Chinese, Indian,
and Brazilian grains researchers and their work are being felt
globally. Different countries also have different types of
involvement of universities in grains breeding, although in
the process of training the next generation of grains breeders,
some universities have developed significant breeding pro-
grams and have become a mainstay of breeding efforts for a
country. Several interdependent factors have given rise to
greater commercial involvement in grains breeding, including
the increased importance of plant breeders rights, develop-
ment and application of biotechnology in grains science,
shrinkage of government support for agricultural research,
and the increased importance of intellectual property protec-
tion. In some cases, privatization of former public sector plant
breeding institutes has taken place.
In some parts of the world, such as Europe, there has been
a long history of significant plant breeding efforts being
carried out by industry; in others, such as Australia (with the
exception of hybrid cereals), almost all plant breeding was
formerly carried out by government and universities. Nowa-
days, Australia and several other countries separate the respon-
sibilities for prebreeding and breeding between public and
private sector organizations, respectively. The other industry
group that has become more involved in grains research has
455
been farmers themselves. Farmer-driven groups now often
have a key role in identifying research priorities as well as
being a vehicle for dissemination of research results in a
number of countries, farmer groups directly manage on-farm
adaptive research trials on technologies such as rotations, weed
control, and reduced tillage, but some are also adopting a
wider perspective. Realizing that both small and large farms
are often diversified enterprises, grains research has a much
greater emphasis on farming systems than in previous years.
The influence of farmers is important in fitting grains research
into a wider farming systems context. Organizations that use
farmers levies to fund research such as the Home Grown
Cereals Authority in the United Kingdom or the Australian
Grains Research and Development Corporation have also
been instrumental in encouraging greater farmer involvement.
Grains science is carried out within formal institutional
structures, most of which have evolved from a range of histor-
ical and political situations, rather than having been developed
with a consideration of whether the institutional structures fit
the needs of the twenty-first century. In many state- or
provincial-based departments, there remains a close collabora-
tive linkage between cereal breeding and cereal chemistry, and
the scientists involved carry out a service function as well as
independent research often creating a tension between the
time spent on the two, but with the advantage that the research
objectives are informed by genuine industry problems.
There are few grains science institutions globally that cover
all of cereals, legumes, and oilseeds. Instead, there may be
research clusters within university agronomy or agriculture
faculties or grains researchers within plant science or crops
research organizations. It is more usual for postharvest tech-
nology institutions or grain storage research laboratories to
consider various issues relating to a wider range of cereals
and other grains together, on the basis that the storage and
handling of grains as durable commodities have common
research challenges. Over the last decade, there has been a
decrease in postharvest grains research globally, given that
there is a prevailing view that the main priority is now adop-
tion of established technologies.
Some institutes do cover several cereals or grains more
broadly, although usually with a focus on those crops relevant
for their country or targeted region. For example, wheat and
barley are often studied together. Some of the equipment and
techniques are common, while others specialized for exam-
ple, while Falling number is used as a measure of postharvest
sprouting for both wheat and barley, dough rheology testing is
specific to wheat. Specialized grains science departments are
found in a small number of US and European universities, but
their numbers are decreasing. Arguments for this model are the
critical mass of researchers and ability of students to train in
aspects from grains breeding to postharvest technology. On the
other hand, such departments can lack linkages to emerging or
discovery science and to the broader range of disciplines in
agriculture and science.
One of the lessons of modern molecular biology is that
techniques such as those in functional genomics are often
readily transferable between grains, other crops, microorgan-
isms, and animals. This aspect has very much been exploited in
recent years as whole genome sequencing becomes faster and
much less expensive, and its use to understand disease
456 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Global Trends and the Commercial Sector
resistance and quality associations becomes more widespread.
The high degree of synteny tells us that gene structure and
function are conserved across cereals and the great similarity
of their physiological and biochemical processes often makes it
simple to utilize research ideas and techniques between differ-
ent grain crops.
There are no definitive answers to the issue of an optimal
institutional structure for grains research other than to assume
that earlier structures need not necessarily be more important
than ever for breeders, agronomists, crop protection scientists,
and researchers of processing quality to interact. Finally, insti-
tutional structures can coevolve with the evolution of technol-
ogy. With the growth of networks of researchers collaborating
nationally and globally and able now to exchange large data
sets through cloud computing, the establishment of formal
institutional structures and a prerequisite for geographic co-
location to foster collaboration has become less important. But
humans will remain social animals, and it is hard to envisage a
world where face-to-face collaboration will become irrelevant.
Exercises for Revision
What are some of the factors leading grains research to be
more demand-driven?
How has greater collaboration between research institutes
been encouraged by governments and funding agencies?
What type of research do grains processing companies in
Europe and North America predominantly carry out?
What opportunities stimulated agrochemical companies to
get involved in grains breeding?
What are the main areas of subject matter emphasis for
commercial grains research and development in India and
Africa?
What factors have led to greater commercial involvement in
grains breeding?
Exercises for Readers to Explore the Topic Further
Explore the similarities and differences between focus areas
for wheat and barley research. Why are some areas similar
yet others quite different?
How do priorities for research relating to irrigated wheat
and rice differ from those for rainfed (or dryland) wheat
and rice?
Compare and contrast the role, organization, and business
models of farmers organizations undertaking grains
research in a developed country and a developing country.
See also: Barley, Rice and Maize Processing: Barley: Malting;
Breeding of Grains: Barley: Genetics and Breeding; Canola/
Rapeseed: Genetics and Breeding; Maize: Breeding; Soybean:
Germplasm, Breeding, and Genetics; Wheat Breeding: Exploiting and
Fixing Genetic Variation by Selection and Evaluation; Food Grains
and the Consumer: Consumer Trends in Grain Consumption;
Genetically Modified Grains and the Consumer; Grains and Health;
Genetics of Grains: Development of Genetically Modified Grains;
Genome Mapping; Genomics of Food Grains; Processing of Grains:
Cereals: Breakfast Cereals; The Basics: The Grain Crops: An
Overview; The Cereal Grains: An Overview of the Family of Cereal
Grains Prominent in World Agriculture; Wheat-Based Foods:
Breads; Cookies, Biscuits and Crackers: Formulation, Processing and
Characteristics; Pasta.
Further Reading
Alston JM, Pardey PG, and Smith V (1999) Paying for International Productivity.
Baltimore/London: Johns Hopkins University Press.
Alston JM, Beddow JM, and Pardey PG (2009) Agricultural Research, productivity and
food prices in the long run. Science 325: 12091210.
Cuesta-Marcos A, Kling JG, Belcher AR, Filichkin T, Fisk SP, Graebner R, et al. (2016)
Barley: genetics and breeding. In: Corke H, Wrigley C, Faubion J, and
Seetharaman K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 4, pp. 287295.
Elsevier.
DePauw R and OBrien L (2016) Wheat Breeding: Exploiting and Fixing Genetic
Variation by Selection and Evaluation. In: Wrigley C, Seetharaman K, Corke H, and
Faubion J (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 4, pp. 279286.
Elsevier Ltd.
Evenson RE and Gollin D (2003) Assessing the impact of the Green Revolution,
19602000. Science 300: 758762.
Henry RJ (2016) Genomics. In: Corke H, Wrigley C, Faubion J, and Seetharaman K
(eds.) Encyclopedia of Food Grains, 2nd edn., vol. 4, pp. 360364. Elsevier Ltd.
Jetter KM and Cassady DL (2006) The availability and cost of healthier food alternatives.
American Journal of Preventative Medicine 30: 3844.
Jones JM and Sheats DB (2016a) Grains and Health: Misinformation and
Misconceptions. In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.)
Encyclopedia of Food Grains, 2nd edn., vol. 2, pp. 2328. Elsevier Ltd.
Jones JM and Sheats DB (2016b) Consumer Trends in Grain Consumption.
In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of Food
Grains, 2nd edn., vol. 2, pp. 2934. Elsevier Ltd.
Pardey RG and Beintema NM (2001) Slow Magic: Agricultural R&D a Century after
Mendel. Washington, DC: International Food Policy Research Institute.
Pray C, Gisselquist D, and Nagarajan L (2011) Private investment in agricultural
research and technology transfer in Africa. Agricultural R&D: Investing in Africas
future, ASTI-FRA Conference, Accra, Ghana, 57 December, Working paper 13.
Pray CE and Nagarajan L (2012) Innovation and Research by Private Agribusiness in
India. Washington, DC: International Food Policy Research Institute IFPRI
Discussion Paper 01181.
Ray DK and Foley JA (2013) Increasing global crop harvest frequency: Recent trends
and future directions. Environmental Research Letters 8: 044041. http://dx.doi.org/
10.1088/1748-9326/8/4/044041 10 pp.
Saltzman A, Birol E, Bouis HE, et al. (2013) Biofortification: Progress toward a more
nourishing future. Global Food Security 2: 917.
Skerritt JH (2016a) Research organizations of the world: Europe and North America.
In: Corke H, Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of Food
Grains, 2nd edn., vol. 1, pp. 436443. Elsevier Ltd.
Skerritt JH (2016b) Research organizations of the world: Asia-Pacific, Central-South
America, and Africa-Middle East. In: Corke H, Wrigley C, Faubion J, and
Seetharaman K (eds.) Encyclopedia of Food Grains, 2nd edn., vol. 1. pp. 444450.
Elsevier Ltd.
Tiwari VK, Faris JD, Friebe B, and Gill BS (2016) Genome mapping. In: Corke H,
Wrigley C, Faubion J, and Seetharaman K (eds.) Encyclopedia of Food Grains,
2nd edn., vol. 4, pp. 365375. Elsevier Ltd.
US Energy Information Administration (2012) Biofuels Issues and Trends, October
2012, US Department of Energy, Washington (www.eia.gov/biofuels/issuestrends/
pdf/bit.pdf).
 Scientific Societies Associated with Grain Science
SC Nelson, AACC International (AACCI), St. Paul, MN, USA
CW Wrigley, The University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Large numbers of scientists and technologists are involved
with food grains worldwide.
These people have formed scientific associations, based on
their specific interests, to facilitate interactions and infor-
mation sharing.
These interactions involve working collaborations and the
exchange of ideas via conferences, publications, standard-
izing analytic methods, and explaining grain science to the
general public.
Two such associations, originally based in the United States
and in Europe (the AACCI (American Association of Cereal
Chemists International) and the ICC (International Associ-
ation for Cereal Science and Technology)), have become
international.
Other associations have regional memberships.
Some of these associations specialize in specific aspects
such as seeds, breeding, agronomy, soil science, and grain-
related health benefits.
Learning Objectives
Achieve understanding of the need for and the activities
of scientific associations that have been established to
facilitate collaboration and exchange of ideas between
scientists and technologists involved in the breeding,
growing, harvesting, marketing, and processing of food
grains and nutritionists and consumer groups.
Introduction
The industries that grow, harvest, and process grain are big
businesses worldwide, involving large numbers of people,
including many scientists and technologists. It is important
that all these professionals should have opportunities to inter-
act with one another, to exchange ideas, to learn from one
another, and to develop grain-testing methods that are
accepted internationally.
In addition, they need to interact more broadly with con-
sumers, the media, and government agencies. Opportunities
for these various activities are provided by specialist societies
and associations (learned societies), with grain scientists as
members, formed for the purposes of interacting with one
another in their areas of expertise.
Many such scientific societies have been formed for grain
scientists, covering the wide range of aspects of grain science,
from breeding, to agronomy and farming, to the chemistry of
the harvested grain, to marketing, processing, and retailing.
Some of these societies are selected for description in the
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00060-7
succeeding text, but the range covered is by no means compre-
hensive. In addition to scientific societies, there are also trade
associations, formed for the purposes of interaction at the
technical and commercial levels and for lobbying purposes. A
list of many scientific and trade associations of the food indus-
try has been assembled by www.stagnitomedia.com and www.
hoovers.com.
The activities of the scientific associations include the orga-
nization of conferences at which lectures are presented, as
shown in Figure 1. In addition to lectures, there are poster
presentations, involving the author standing beside the
description of a research project on a large poster board for
discussion with conference delegates (Figure 2).
Conferences also serve the purpose of providing opportu-
nities for personal interactions to occur; these face-to-face sit-
uations complement the less personal interactions by phone
and Internet at other times. Trade exhibits (Figure 3) at con-
ferences provide information about the latest equipment
and resources. These learned societies use conference time for
making awards to their members who have excelled in aspects
of the science and technology relevant to the societys activities.
Figure 4 shows the recent presentation of several awards
at a meeting of AACCI. The Thomas Burr Osborne Medal
(Figure 5) is one of these awards.
Other activities include the dissemination of information
via journals, books, and other publications; the establishment
of professional standards of conduct and training; and the
development of standard methods of analysis.
A related activity is the provision of check samples
characterized subsamples, such as grain or flour, that are pro-
vided to many cooperating laboratories for them to establish
that standard methods of analysis are providing similar results
across various regions.
The provision of reliable scientific information is becoming
an important function, thereby advising decision-making bod-
ies and the general public; these activities may even border on
lobbying when it becomes apparent that misinformation is
prevailing.
The Range of Societies
Prominent among these learned societies are those relevant to
the quality and processing of grain. In particular, AACCI has a
worldwide membership of scientists involved in grain produc-
tion and processing. The ICC, based in Vienna, Austria, is an
international association committed to international coopera-
tion through disseminating information and developing stan-
dard methods relevant to grain production and processing.
There are also national bodies with similar objectives, such as
the Chinese Cereals and Oils Association (CCOA), based in
Beijing, China, and the Australasian Grain Science Association
457
458 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Scientific Societies Associated with Grain Science
Figure 1 A major activity of scientific associations is the organization of
conferences at which lectures are presented to inform members about
recent developments in their field of grain science.
Figure 2 Poster sessions provide an alternative to lectures at
conferences. The author of the information on the poster is available to
discuss its contents with other delegates.
Figure 3 Trade exhibits at conferences help members become familiar
with the latest equipment and resources.
(previously the Cereal Chemistry Division of the Royal Austra-
lian Chemical Institute (RACI)).
Other societies have been formed for those involved in
specific aspects of chemical analysis or individual classes of
chemical compounds. For example, the American Oil Chem-
ists Society (AOCS), comprising some 5000 members
Figure 4 Scientific societies honor members who excel in their field of
study by the presentation of awards.
Figure 5 The Thomas Burr Osborne Medal, awarded for research in
the field of cereal chemistry that has contributed significantly to
the progress of the science. The medal commemorates an important
research pioneer of cereal chemistry early in the twentieth century.
worldwide, provides a global forum for the science and tech-
nology of fats, oils, surfactants, and related materials. Further-
more, AOAC International is an internationally recognized
organization with 130 years of experience in validating and
approving analytic methods for foods and agriculture. In these
cases, involvement goes well beyond the analysis of grains, but
grains are a significant part of their broader scope.
The American Society of Agronomy, the Crop Science Soci-
ety of America, and the Soil Science Society of America attract
scientists involved in plant growth and production. While their
interests include grain production, the involvement of these
societies covers the full breadth of plant-based agriculture.
Complementary to this aspect of grain science is the Interna-
tional Seed Testing Association (ISTA), concerned with the
earliest stage of plant production, namely, the provision of
pure seed. Again, the scope of this society extends well beyond
food grains to cover all types of plant seeds.
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Scientific Societies Associated with Grain Science
AACC International (AACCI)
AACCI is one of the largest international nonprofit organiza-
tions dedicated to creating opportunities for providing and
exchanging knowledge and advancing innovations across the
broad disciplines of cereal grain science by facilitating research,
education, collaboration, technical service, and advocacy efforts.
AACCI was founded in 1915 for the purpose of standardizing
methods of analysis among cereal laboratories. Today, the
associations mission is to be the global authority in cereal grain
science and technology. The association now provides a compre-
hensive and respected collection of grain science methods. All
methods go through a rigorous approval process by an AACCI
technical committee. AACCIs technical committees are com-
posed of scientists chosen for their specific areas of expertise.
Today, as industry issues arise, AACCI strives to provide
science-based approaches and responses to address current
needs and trends. The diverse membership provides the creative
and scientific strength to help provide creative solutions and
quality recommendations in reaction to a dynamic industry
and world environment. Through its activities in methods val-
idation, AACCI is acknowledged as a scientific authority when
proposing definitions and providing comments to regulatory
agencies worldwide. Key definitions developed include dietary
fiber, whole grain, oat bran, glycemic carbohydrates, and aleu-
rone. Additionally, AACCI has provided comments on whole
grain label statements; proposed guidance on food allergens,
food labeling, and nutrition with respect to gluten-free foods;
and health claims relating to the consumption of certain oat
products to a reduced risk of coronary heart disease. In 2013,
AACCIs Whole Grains Working Group approved its character-
ization of a whole grain. The characterization states that a
whole grain food product must contain 8 g or more of whole
grain per 30 g of product. Volunteers have also organized ad
hoc internal committees to examine key issues affecting the
science and developed action plans to address these issues.
AACCIs strategic initiatives include (1) generating and dis-
seminating the highest quality of science and information on
cereal grains and their products; (2) driving scientific commu-
nication, collaboration, and innovation; (3) providing profes-
sional development for its members; (4) facilitating the
interface of the many disciplines involved in cereal and grain
science; and (5) ensuring a vital organization in future years. In
2015, AACCI commemorated its centennial year by examining
the major advances made in cereal grain science through the
associations 100 years. AACCI produced a series of feature
articles in its publication, Cereal Foods World, examining the
advances by category and explaining the importance of these
advances in science today. Centennial activities also included
the creation of a timeline of historic advances and key moments
in AACCI history, which is currently available on AACCIs
website. AACCIs 2015 Centennial Meeting included additional
scientific content and a speaker who looked to how todays
understanding of cereal grain science will inform the future of
the science and its impact on the world.
The Activities of AACCI Members
Many AACCI members are involved in food manufacturing
using grains, one of the fastest-growing, most dynamic
459
segments of the food industry. They are responsible for pro-
ducing some of the most popular and profitable products, such
as new types of cracker, cookies, pretzels, or other snack foods.
Bringing such products into the market involves design, devel-
opment, and marketing skills. Breads, tortillas, frozen doughs,
and dry mixes all rely on cereal chemists for their success in the
marketplace.
Other AACCI members are involved in the upstream of the
food industry. They may be associated with plant-breeding
programs, with responsibility for ensuring that the grain from
new varieties is suited for specific marketing and processing
requirements. Research activities are prominent for other cereal
chemists, with the objectives of elucidating the chemical and
molecular bases of grain quality, thereby laying a foundation
for the development of improved testing methods for grain
quality. Other research activities are likely to lead to the devel-
opment of new grain types with better processing and nutri-
tional attributes.
AACCI Resources
AACCI has provided resources in grain science for a century,
supporting members professional needs in private industry,
academia, and government. The association provides an ave-
nue of bringing together and disseminating scientific informa-
tion and technical research on cereal grains and related
materials and their processing and utilization. This informa-
tion is made available through annual meetings, through con-
tinuing education courses, and in publications. The range of
published materials includes scientific journals, books, elec-
tronic newsletters, and data provided on the website.
AACCI publishes Cereal Chemistry, a journal with peer-
reviewed, original research on raw materials, processes, and
products that relate to the utilization of cereal grains, oilseeds,
and pulses and analytic procedures and methods in the grains
area. AACCI also publishes Cereal Foods World, which includes
feature articles and original research that focus on advances in
grain-based food science and the application of these advances
on current practices in baking, snack foods, breakfast foods,
and other grain-based products. In addition to these two jour-
nals, AACCI has published more than 80 titles on various food-
science topics. Some titles are technically focused while others
are more general. The AACCI handbook series offers a single
source of practical information for the major ingredients used
in food processing.
AACCIs website offers members the opportunity to obtain
information and resources in one common location. The web-
site features an online catalog of books, special reports, mem-
bership directory, a calendar of events, and online symposia.
AACCI journals and electronic copies of publications can be
found in the AACCI online Grain Science Library. This content
platform delivers a large and growing collection of resources,
including Cereal Chemistry, Cereal Foods World, and AACCI
PRESS online books. AACCIs continuing education programs
offer professional development services for food industry pro-
fessionals at any level in a variety of food-related industries.
Core services include short courses taught by experts that offer
comprehensive training and tools for real-world applications.
AACCIs continuing education programs also focus on emerg-
ing issues and strategies for the future success in the food
460 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Scientific Societies Associated with Grain Science
industry. Members receive special rates on technically focused,
hands-on training. Courses are offered globally.
AACCI Meetings
AACCIs annual meetings are attended by an international
audience of more than 1000 grain-based professionals, provid-
ing a rich scientific program and networking opportunities for
the global grain industry. Presenters have the opportunity to
share the latest research on topics such as biotechnology, func-
tional foods and ingredients, health and nutrition, structure
and function of food components, and quality enhancements.
The annual meeting also provides students with numerous
opportunities to display their research, including product
development and research paper competitions. The annual
meeting features a trade show, typically composed of approx-
imately 250 exhibits.
AACCI recognizes excellence through its awards program
(Figure 4). AACCI awards include the AlsbergFrenchSchoch
Memorial Lectureship, the C. W. Brabender Award, the Edith A.
Christensen Award, the Excellence in Teaching Award, AACCI
Fellowships, the Phil Williams Applied Research Award, the
Texture Technologies Quality Research Awards, the Thomas
Burr Osborne Medal (Figure 5), the William F. Geddes Memo-
rial Award, the Young Scientist Research Award, and Honorary
Memberships. In addition, the AACCI Foundation provides
scholarships and fellowships annually for students majoring
in disciplines related to cereal science. Student and young pro-
fessionals receive travel assistance grants to attend the annual
meeting through the Student Travel Award program and the
Cecil F. Pinney Travel Award.
Members have the opportunity to attend local section meet-
ings, connecting them to other professionals within their own
region. AACCI also has several subject-matter divisions focused
on basic research and development areas. The Student Associ-
ation, in conjunction with the Professional Development
Panel, provides students with professional development
opportunities such as showcasing their research (with lectures
and poster presentations; Figures 1 and 2), interacting with
well-respected members of the scientific community through
mentorships, and building professional leadership skills
through the competitions mentioned previously. Students
also have opportunities to attend hands-on workshops includ-
ing the AACCI Breakfast Cereals and Europe Sections Euro-
pean Young Cereal Scientists and Technologists Workshop and
the Young Cereal Chemists Meeting.
The International Association for Cereal Science
and Technology
The ICC is an independent, internationally recognized organi-
zation of experts specializing in the milling of wheat and other
cereals, bread making, and the production of other cereal-
based foods from around the world. In more recent times,
the ICC has expanded its focus to address issues that contribute
to improved food quality, food safety, and food security for the
health and well-being of all people. The ICC is
an apolitical forum for all cereal scientists and
technologists;
a publisher of international standard methods and a scien-
tific journal;
an organizer of major national and international events in
the field;
a promoter of international cooperation on a global,
regional, and national level;
a significant player in the coordination and participation in
international research projects.
Objectives
The ICC is a nonpolitical, nonprofit-making organization.
Membership is open to all interested countries, corporations,
and institutions that are prepared to offer their cooperation
and to support the aims and ideals of the association.
A Short History of the ICC
The ICC was founded in 1955 on the occasion of the third
International Bread Congress in Hamburg, Germany, as the
International Association for Cereal Chemistry (ICC).
The original objective of the ICC was the development of
internationally approved and accepted standard testing pro-
cedures for cereals and flour. Today, the ICC is one of the
foremost international organizations in the field of grain sci-
ence dedicated to international cooperation, the dissemination
of knowledge, and the improvement in safety and quality of
cereal-based foods. The association has its headquarters and its
General Secretariat in Vienna, Austria. At present, some 30
countries from five continents are represented in the ICC.
ICC Mission Statement
ICC is the preeminent international association in the field of
cereal science and technology, committed to international
cooperation through the dissemination of knowledge, con-
ducting research, and developing standard methods that con-
tribute to advance innovation and improve food quality, food
safety, and food security for the health and well-being of all
people.
National Cereal Chemistry Societies
Many countries have formal and informal associations of pro-
fessionals in grain science. In addition, international associa-
tions, such as the AACCI, have regional sections and branches
that provide opportunities for members to hold local
meetings.
The Chinese Cereals and Oils Association
The CCOA, based in Beijing, China, provides opportunities for
interaction for a large number of Chinese scientists working
with grains, edible oils, and related commodities. While its
activities are mainly centered within China, the association
also hosts international conferences, thereby involving the
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Scientific Societies Associated with Grain Science
wider range of grain scientists. The CCOA has over 6000 indi-
vidual members and has over 570 organization members. The
CCOA is a member of the ICC. The CCOA is divided into four
subassociations, namely, grain storage, grain food, edible-oil
processing, and animal-feed production. The CCOA publishes
a journal, the Journal of the Chinese Cereals and Oils Association.
The Chinese version is published every 2 months, while the
English version appears annually.
Australian and New Zealand Grain Scientists
The Australasian Grain Science Society (AGSA) is currently the
main association for cereal chemists and other grain scientists
in Australia, New Zealand, and nearby regions. AGSA, now a
stand-alone society, has grown out of the Cereal Chemistry
Division, which was a part of the RACI, based in Melbourne,
Australia. In addition, many Australian grain scientists are
members of the AACCIs Downunder Section.
AGSA provides opportunities for scientists and technolo-
gists to exchange information and to interact with colleagues
internationally, especially at annual meetings. These annual
meetings continue the series of annual Australian Cereal
Chemistry Conferences, which commenced in 1950, so that
the 2016 annual meeting is the 66th conference in this series.
At regular intervals, these meetings are combined with other
international societies, especially the AACCI and the ICC.
Other activities include workshops and training seminars
on specific aspects of grain science, the provision of standard
methods of analysis, and facilities for checking samples with
known analytic results by which laboratories are able to com-
pare the results of their analytic methods with others.
AGSA has several awards to acknowledge special contribu-
tions to grain science. The Guthrie Medal (Figure 6)
Figure 6 The Guthrie Medal, originally awarded by the Cereal Chemistry
Division of the Royal Australian Chemical Institute, is now awarded of the
Australasian Grain Science Association. It is awarded for outstanding
contributions in the field of grain science. The medal commemorates an
Australian cereal chemist of the 1890s and 1900s.
461
commemorates the pioneering wheat research of Australian
Frederick Bickell Guthrie, who formed the worlds first
chemistbreeder collaboration with the wheat breeder William
Farrer commencing in the 1890s.
Throughout the life of AGSA and the Cereal Chemistry
Division, presentations at annual meetings have been recorded
in published form, in recent years under the titles Cereals 2016,
Cereals 2015, Cereals 2014, etc. Details of these publications
and other activities are provided on the AGSA website (http://
ausgrainscience.org.au).
Societies Involved with Grain Growth
The International Seed Testing Association
The ISTA is a worldwide, nonprofit association, whose main
activity is to provide methods and services for the testing of
seed traded internationally. The primary purpose of the
association is thus to develop, adopt, and publicize standard
methods for sampling and testing seeds of all types, not only
the edible grains. ISTAs secondary purpose is to promote all
areas of seed science and technology. These purposes are
largely served by the publication of relevant handbooks and
bulletins, scientific journals (Seed Science and Technology
and Seed Symposium Abstracts), proceedings of symposia and
workshops, and training booklets.
The primary ISTA instrument for promoting uniformity of
seed-testing procedures is the International Rules for Seed Test-
ing, a publication that is updated annually with amendments
and additions resulting from ISTA meetings. The ISTA Secre-
tariat is based in Bassersdorf, Switzerland.
The American Societies of Agronomy, Crop, and Soil Science
Founded in 1907, the American Society of Agronomy is
dedicated to the development of agriculture enabled by sci-
ence, in harmony with environmental and human values. The
society supports scientific, educational, and professional activ-
ities to enhance communication and technology transfer
among agronomists and those in related disciplines on topics
of local, regional, national, and international significance.
The Crop Science Society of America is an educational and
scientific organization composed of more than 5000 members
dedicated to the advancement of crop science. The motto is
Plant science for a better world. Founded in 1955, the society
is international in scope with members in more than 100
countries who are advancing the discipline of crop science by
acquiring and disseminating information about crops in rela-
tion to genetics and plant breeding, crop physiology and pro-
duction, germplasm resources, and environmental quality. An
important function of the society is the production of its main
journal Crop Science.
The Soil Science Society of America, established in 1936, is
the professional home for over 6000 professionals throughout
the world involved in soil science. Their claim is Soils sustain
life. The primary purpose of the society is to advance the
discipline and practice of soil science by study and education
concerning soils in relation to crop production, environmental
quality, ecosystem sustainability, bioremediation, waste man-
agement and recycling, and wise land use.
462 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Scientific Societies Associated with Grain Science
Societies Involved with Grain Analysis
AOAC International
AOAC International is a globally recognized, independent,
third party, not-for-profit association with 130 years of expe-
rience in validating and approving analytic methods for foods
and agriculture. It has over 3000 members worldwide.
The organization was founded in 1884 as the Association of
Official Agricultural Chemists, under the auspices of the US
Department of Agriculture. Its initial role was to adopt uniform
methods of analysis for fertilizers. In the following year, a
convention established the AOAC as an independent organi-
zation, but membership was restricted to analytic chemists in
state and federal positions of the US government, and this
membership requirement remained for a century. In 1965, to
recognize the expansion of AOACs scope of interest beyond
agricultural topics, the associations name was changed to the
Association of Official Analytical Chemists.
During the 1970s, membership was extended to scientists
outside the United States and to nongovernmental scientists,
with many members working in industrial laboratories. Dur-
ing the 1980s and 1990s, the scope of analyses expanded and
there was increasing demand for quality control of laboratories
and international laboratory accreditation. Consequently, the
name of the association was changed to AOAC International,
thereby retaining the initials by which the association had been
known for 130 years, while eliminating reference to a specific
scientific discipline or profession.
The scope of the AOAC includes methods not only for the
analysis of grains but also for a much wider range of materials.
AOAC provides a number of key publications (especially its
Official Methods of Analysis), hosts technical meetings and con-
ferences, and offers training courses in the areas of laboratory
management, quality assurance, accreditation, statistics, and
measurement uncertainty.
The American Oil Chemists Society
The AOCS (Your Global Fats and Oils Connection) is another
American scientific association that has become international,
growing and expanding in scope since its origins in 1915. This
scope now includes oil- and fat-related commodities, oilseeds,
oilseed meals, and edible fats. Grains are thus prominent in
this range of materials for analysis, but the scope goes beyond
grains, including other sources of edible fats, such as fish and
animals. The AOCS technical services are based in Champaign,
IL, the USA.
Societies Relating to Food Grains and Health
Grains for Health Foundation
The people working with Grains for Health Foundation
include scientists, business leaders, educators, and health pro-
fessionals working together at the grassroots level encouraging
all sectors, disciplines, and communities to use their expertise
to improve public health.
The Grains for Health Foundation provides publications,
resources, and events within the grains and health
communities. Membership helps shape this foundations
objectives and foster integral conversations that create practical
change. The Grains for Health Foundation actively seeks both
sponsor members and partner members.
Grains for Health Foundation goals include redesigning the
food supply by facilitating the development, delivery, and
consumption of grain-based foods in order to promote a
healthy body weight, reduce chronic disease, and curb health-
care costs. These bridges ignite the necessary conversations
around making healthy, grain-based foods accessible.
Grains for Health Foundation measures its success to the
extent that contributions help build and maintain the crucial
relationships needed between the whole grain food supply and
the researchers and policy makers that regulate the distribution
and education surrounding grain-based foods. Measures of
success include the following:
Foods with healthy grains are available in your grocery,
school, or business.
Health-care professionals recommend whole grains as a
means of reducing chronic illness.
Policy makers design legislation leading to healthier grain
products.
Scientists create grains able to feed our communities.
Rates of chronic diseases decline.
Cost of health care is lowered.
Grains for Health Foundation undertake strategic and transfor-
mative work on all levels of the grain chain from connecting
growers to scientists to politicians to distributors to educators
to health-care professionals. The cross-disciplined work builds
a vast community needed to create the positive change in
public health so you can be assured access to healthy, grain-
based foods.
The HEALTH GRAIN Forum
This association was founded in May 2010 to continue promo-
tion of science and communication for production and con-
sumption of healthy cereal foods, as a follow-up of the
HEALTHGRAIN Forum of the European Union. There are cur-
rently members from 19 countries. The organizational struc-
ture includes a board and four task forces to plan activities,
interacting via telemeetings on a regular basis. Two annual
workshops are organized to provide opportunities for
interaction.
The mission of the HEALTHGRAIN Forum is to promote
science-based concepts fully, unlocking the health-promoting
potential in the entire grain food production chain to obtain
healthy, convenient, and appealing foods. The vision is to
assist consumers in health maintenance worldwide, help
reduce health-care costs, and provide added value for compa-
nies in the production chain. The motto is The best of the grain
for consumer well-being.
Celiac Societies
Celiac societies have been formed in many countries and
regions by people with celiac disease and related forms of
dietary intolerance to gluten. These are not associations of
 SCIENTIFIC ORGANIZATIONS RELATED TO GRAINS | Scientific Societies Associated with Grain Science
professionals only, but rather of people with celiac disease,
together with their relatives and friends. They may, in turn,
rely on scientists and medical workers to provide advice in the
forms of publications, consultation, and conference presenta-
tions for the benefit of the members. A few web addresses are
provided in the list in the succeeding text, but there are many
more such societies worldwide.
Future Prospects
Into the future, grain science professionals will continue to
need opportunities to interact and collaborate. Scientific soci-
eties can continue to provide scientists not only with validated
and archived information but also with the essential feature of
contact with other scientists. While todays technology pro-
vides new options for frequent and instant communications,
it continues to be important for scientists to meet face to face
for effective dialogue. Some means of interaction will progres-
sively change, largely due to changes in the range of novel
electronic media. Already, traditional means of print publica-
tion are being duplicated or replaced by various electronic
means. The value of face-to-face meetings and conferences
cannot be readily replaced.
Exercises for Revision
What features distinguish the associations described in this
article from other types of associations in society in general?
Which of the associations described would be the most
appropriate for you, depending on your location and your
area of grain science (e.g., breeding, farming, milling, food
processing, and advising about nutrition)?
List the range of activities and opportunities provided by
membership of a scientific society that would not be other-
wise available.
Exercises for Readers to Explore the Topic Further
There are even more associations relevant to grain science
than the prominent examples given in this article. What
463
other examples can you find? Think, for example, of asso-
ciations involving breeders, geneticists, grain growers,
health professionals, nutritionists, and consumers.
Find out the costs of membership, of conference atten-
dance, and of other forms of involvement for some of
these associations. Try to assess the value for money in
terms of professional benefits to be gained by members.
To what extent do you reckon that benefits depend on
involvement via publications (only), conference atten-
dance (only), and/or volunteering for direct participation
in association affairs?
Look for indications that these associations have recently
been involved in advising decision-making bodies with
respect to their areas of expertise.
See also: Food Grains: Intolerance, Allergy and Diseases:
Celiac Disease; Grain Composition and Analysis: Standardized
Test Methods for Grains and Grain-Based Products; Scientific
Organizations Related to Grains: Research Organizations of the
World: Asia-Pacific, CentralSouth America, and AfricaMiddle East;
Research Organizations of the World: CGIAR; Research Organizations
of the World: Europe and North America; Research Organizations of the
World: Global Trends and the Commercial Sector.
Relevant Websites
http://www.aaccnet.org The AACC International.
http://www.agronomy.org The American Society of Agronomy.
http://www.aoac.org AOAC International.
http://www.aocs.org The American Oil Chemists Society.
http://www.asbcnet.org The American Society of Brewing Chemists.
http://www.ccoaonline.com Chinese Cereals and Oils Association.
http://www.coeliac.org.au The Coeliac Australia.
http://www.crops.org The Crop Science Society of America.
http://www.csaceliacs.org The Celiac Support Association, USA.
http://www.icc.or.at The International Association for Cereal Science and Technology
(ICC).
http://seedtest.org The International Seed Testing Association.
http://www.nopa.org The National Oilseed Processors Association.
http://www.soils.org The Soil Science Society of America.
http://www.spcouncil.org The Soy Protein Council.
ENCYCLOPEDIA OF
FOOD GRAINS
Second Edition
This page intentionally left blank
 ENCYCLOPEDIA OF
FOOD GRAINS
Second Edition
VOLUME 2
NUTRITION AND FOOD GRAINS
EDITORS
COLIN WRIGLEY
University of Queensland, Brisbane, Australia

HAROLD CORKE
University of Hong Kong, Hong Kong, Peoples Republic of China

KOUSHIK SEETHARAMAN{

JON FAUBION
Kansas State University, Manhattan KS, USA

{
deceased

AMSTERDAM BOSTON HEIDELBERG LONDON

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First edition 2004 (as Encyclopedia of Grain Science)

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CONTENTS

Preface xvii
Tributes to Former Co-Editors xix
Editors Biography xxiii
Editorial Advisory Board xxv
Contributors xxvii

VOLUME 1

THE WORLD OF FOOD GRAINS 1

The Basics 1
The Grains that Feed the World 1
CW Wrigley, H Corke, and J Faubion

Course Structures: Based on EFG Articles 13

H Corke

The Grain Crops: An Overview 16

RA Graybosch

The Grain Chain: The Route from Genes to Grain-Based Products 22

CW Wrigley

Taxonomic Classification of Grain Species 31

LA Morrison and CW Wrigley

Grain: Morphology of Internal Structure 41

H Corke

Grain and Plant Morphology of Cereals and how characters can be used to identify varieties 51
J Wyatt

The Cereal Grains 73

An Overview of the Family of Cereal Grains Prominent in World Agriculture 73
CW Wrigley

Cereals: Domestication of the Cereal Grains 86

LA Morrison

Maize: Overview 99
MP Scott and M Emery

Wheat: An Overview of the Grain That Provides Our Daily Bread 105
CW Wrigley

v
vi Contents

Durum Wheat: Overview 117

GP Kadkol and M Sissons

Rice: Overview 125

BO Juliano

Wildrice, Zizania: Overview 130

EA Oelke and RA Porter

African Rice (Oryza glaberrima): A Brief History and Its Growing Importance in Current
Rice Breeding Efforts 140
JT Manful and S Graham-Acquaah

Barley: An Overview of a Versatile Cereal Grain with Many Food and Feed Uses 147
AM Stanca, A Gianinetti, F Rizza, and V Terzi

Sorghum Grain, Its Production and Uses: Overview 153

A Cruickshank

Rye Grain: Its Genetics, Production, and Utilization 159

HD Sapirstein and W Bushuk

Triticale: Overview 168

BJ Furman

Oats: Overview 173

PK Zwer

Coix: Overview 184

H Corke, Y Huang, and JS Li

Millet Pearl: Overview 190

JRN Taylor

Millet Minor: Overview 199

GK Chandi and GA Annor

Teff: Overview 209

G Bultosa

The Oilseeds 221

Oilseeds: Overview 221
RJ Mailer

Soybean: Overview 228

K Liu

Canola: Overview 237

VJ Barthet

Cottonseed: Overview 242

E Hernandez

Sunflower: Overview 247

GJ Seiler and TJ Gulya

Overview of the Oilseed Safflower (Carthamus tinctorius L.) 254

C Hall III

Linseed: Overview 259

S Cloutier

The Legumes and Pseudocereals 265

Grain Legumes and Their Dietary Impact: Overview 265
TE Michaels
 Contents vii

Pseudocereals: Overview 274

RJ Fletcher

Lupin: Overview 280

DS Petterson

Amaranth: Overview 287

H Corke, YZ Cai, and HX Wu

Beans: Overview 297

SK Sathe

Buckwheat: Overview 307

YZ Cai, H Corke, D Wang, and WD Li

Chickpea: Overview 316

EJ Knights and KB Hobson

Pea: Overview 324

TN Khan, A Meldrum, and JS Croser

Peanuts: Overview 334

RCN Rachaputi and G Wright

Quinoa: Overview 341

SA Valencia-Chamorro

Grains Around the World 349

Grain Production and Consumption: Overview 349
T Beta and C Isaak

Grain Production and Consumption: Africa 359

JRN Taylor

Production and Consumption of Grains: India 367

BS Khatkar, N Chaudhary, and P Dangi

Grain Production and Consumption: China and South-East Asia 374

H Corke and YZ Cai

Grain Production and Consumption: Europe 383

MG Lindhauer

Grain Production and Consumption: Cereal Grains in North America 391

JA Fox and LL Ward

Grain Production and Consumption: Oilseeds in North America 401

PBE McVetty, OM Lukow, LM Hall, I Rajcan, and H Rahman

Grain Production and Consumption in Oceania: Australia and Pacific Countries 409
CW Wrigley and RJ French

Grain Production and Consumption: South America 420

OR Larroque and JC Catullo

Scientific Organizations Related to Grains 429

Research Organizations of the World: CGIAR 429
JH Skerritt

Research Organizations of the World: Europe and North America 436

JH Skerritt

Research Organizations of the World: Asia-Pacific, CentralSouth America, and AfricaMiddle East 444
JH Skerritt
viii Contents

Research Organizations of the World: Global Trends and the Commercial Sector 451
JH Skerritt

Scientific Societies Associated with Grain Science 457

SC Nelson and CW Wrigley

VOLUME 2

NUTRITION AND FOOD GRAINS 1

Food Grains and Well-Being 1

Functional Foods: Overview 1
G Bultosa

Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics 11

G Bultosa

Nutrition: Soy-Based Foods 17

AM Fehily

Food Grains and the Consumer 23

Grains and Health: Misinformation and Misconceptions 23
DB Sheats and JM Jones

Consumer Trends in Grain Consumption 29

JM Jones and DB Sheats

Cultural Differences in Processing and Consumption 35

AMR Hayes and JM Jones

Fortification of Grain-Based Foods 43

CM Rosell

Genetically Modified Grains and the Consumer 50

A Mathiowetz and JM Jones

Labelling of Grain-Based Foods 57

JM Jones

Grains and Health 63

R Korczak, D Hauge, B Maschoff, L Marquart, P Jacques, R Lindberg, and R Menon

Food Grains: Intolerance, Allergy and Diseases 72

Cereal Allergens 72
AS Tatham

Nutrition: Beriberi, A Deficiency Related to Grains 77

KJ Carpenter

Celiac Disease 83
P Koehler, H Wieser, and KA Scherf

The Gluten-Free Diet 91

V Zevallos and I Herencia

Proteins 98
The Protein Chemistry of Cereal Grains 98
F Bekes and CW Wrigley

The Protein Chemistry of Dicotyledonous Grains 109

PR Shewry
 Contents ix

Protein Synthesis and Deposition 115

PR Shewry and P Tosi

Nitrogen Metabolism 123

CA Atkins

Nitrogen in Grain Production Systems 129

XM Zhou, BL Ma, and DL Smith

The Enzymes Associated with Food Grains and Their Functions in Processing 138
DJ Mares, K Mrva, and GB Fincher

Proteomic Analysis Relevant to Cereal Grains 146

A Juhasz, R Haraszi, F Bekes, DJ Skylas, and CW Wrigley

Thiolomics of the Gluten Protein Network of Wheat Dough 154

F Bonomi and S Iametti

Carbohydrates 161
Carbohydrate Metabolism 161
RN Chibbar, S Jaiswal, M Gangola, and M Baga

Starch: Chemistry 174

FL Stoddard

Starch: Synthesis 181

A Regina, S Rahman, Z Li, and MK Morell

Starch: Starch Architecture and Structure 190

V Vamadevan and Q Liu

Starch: Analysis of Quality 198

A Gunaratne and H Corke

Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 208

GB Fincher

Grains other than Cereals: Non-starch Polysaccharides 224

L Ramsden

Resistant Starch and Health 230

A Evans

Health Effects of b-Glucans Found in Cereals 236

SM Tosh and S Shea Miller

Glycemic Index, Glycemic Load, and Their Health Benefits 241

D Dan Ramdath

Fats 248
Lipid Chemistry 248
L Day

Healthy Fats and Oils 257

SM Ghazani and AG Marangoni

Bioactives and Toxins 268

Bioactive Compounds in Wheat Bran 268
HD Sapirstein

Bioactives: Antioxidants 277

T Beta and KG Duodu
x Contents

The Antinutritional Components of Grains 283

CK Madsen and H Brinch-Pedersen

Mycotoxins 290
MT Fletcher and BJ Blaney

Grain Composition and Analysis 297

The Composition of Food Grains and Grain-Based Products 297
CW Wrigley

Standardized Test Methods for Grains and Grain-Based Products 300

AR Bridges and CW Wrigley

Units of Grain Science and Trade: Equivalence between the US, Chinese, and Metric Units 308
W Huang and CW Wrigley

VOLUME 3

GRAIN-BASED PRODUCTS AND THEIR PROCESSING 1

Wheat-Based Foods 1
Cereals: Overview of Uses: Accent on Wheat Grain 1
CF Morris

Breads 8
TR Moore

Flatbreads of the World 19

KJ Quail

Sour Dough Technology 25

SP Cauvain

Cakes, Muffins and Bagels 30

DE Ortiz

Cookies, Biscuits and Crackers: Formulation, Processing and Characteristics 37

SP Cauvain

Cookies: A Diverse Family of Baked Goods 44

S Zydenbos, V Humphrey-Taylor, and CW Wrigley

Wafers: Methods of Manufacture 51

K Tiefenbacher and CW Wrigley

Cookies and Crackers: Commercial Production 59

LC Paulson and CW Wrigley

Noodles: Starch 64
LS Collado and H Corke

Noodles: Asian Wheat Flour Noodles 72

GB Crosbie and AS Ross

Pasta 79
M Sissons

Tortillas 90
LW Rooney and SO Serna-Saldivar

Non-wheat Foods 97
Maize: Foods from Maize 97
SO Serna-Saldivar
 Contents xi

Rice: Chinese Food Uses 110

S Lu

Sorghum: Utilization 116

RD Waniska, LW Rooney, and CM McDonough

Soybean: Soy-Based Fermented Foods 124

DK OToole

Soybean: Soymilk, Tofu, and Okara 134

DK OToole

Food-quality Testing 144

The Application of Sensory Science to the Evaluation of Grain-Based Foods 144
LM Duizer and SB Walker

Noodles: Testing for Quality 154

VA Solah and GB Crosbie

Pasta: Quality Testing Methods 161

A Marti, MG DEgidio, and MA Pagani

Rice: Eating Quality 166

JS Bao

Beverages from Grains 176

Fermentation: Origins and Applications 176
BJB Wood

Fermentation: Foods and Nonalcoholic Beverages 183

JRN Taylor

Beverages: Distilled 193

GH Palmer

Beverages: Asian Alcoholic Beverages 206

DK OToole

Non-food Products from Grains 215

Cereal Grains as Animal Feed 215
JL Black

Pet Foods 223

RCE Guy

Utilization of the Whole Cereal Plant to Maximize its Value 228

L Munck

Fuel Alcohol Production 235

YN Guragain, KV Probst, and PV Vadlani

Biodiesel 245
Y Yan

Processing of Grains 251

Evaluation of Wheat-Grain Quality Attributes 251
CF Morris

Grain Quality Attributes for Cereals Other than Wheat 257

CF Morris
xii Contents

Cereals: Breakfast Cereals 262

EF Caldwell, JD McKeehen, and RS Kadan

Extrusion Technologies 268

RCE Guy

Starch: Uses of Native Starch 274

JW Lawton

Starch: Modification 282

JN BeMiller

b-Glucans: Measurement and Processing 287

MS Izydorczyk

Wheat Processing 299

Milling and Baking: History 299
CE Walker and WD Eustace

Wheat: Dry Milling 307

AK Sarkar and JE Dexter

Wet Milling of Wheat 320

R Velicogna and S Shea Miller

Oven Technologies 325

CE Walker

Bakeries: The Source of Our Unique Wheat-Based Food, Bread 335

JE Bock, CW Wrigley, and CE Walker

Analysis of Dough Rheology in Breadmaking 343

B Dobraszczyk

Frozen Dough 354

D Domingues and C Dowd

Refrigerated Dough 359

D Domingues, C Dowd, and W Atwell

Chemistry of Cake Manufacturing 367

TS Palav

The Gluten Proteins of the Wheat Grain in Relation to Flour Quality 375
F Bekes, MC Gianibelli, and CW Wrigley

Ultrastructure of the Wheat Grain, Flour, and Dough 384

S Grundas and CW Wrigley

Cereal Food Production with Low Salt 396

L Day

Baked Product Staling: Mechanisms, Determinations, and Anti-staling Strategies 403

A Goldstein and K Seetharaman

Gluten and Modified Gluten 408

IL Batey and W Huang

Snack Foods: Processing 414

MN Riaz

Barley, Rice and Maize Processing 423

Barley: Malting 423
L MacLeod and E Evans

Barley: Milling and Processing 434

MS Izydorczyk and JE Dexter
 Contents xiii

Rice Processing: Beyond the Farm Gate 446

L Pallas

Oil from Rice and Maize 453

JS Godber

Maize: Dry Milling 458

KD Rausch and SR Eckhoff

Maize: Wet Milling 467

KD Rausch and SR Eckhoff

Oilseed and Legume Processing 482

Soybean: Soy Concentrates and Isolates 482
C-Y Ma

Soybean: Processing 489

T Wang

Canola: Processing 497

JK Daun and EH Unger

VOLUME 4

THE PRODUCTION AND GENETICS OF FOOD GRAINS 1

Grain Marketing and Grading 1

Wheat: Grading and Segregation 1
RL Cracknell and RM Williams

Wheat: Marketing 9
RL Cracknell and RM Williams

Barley: Grading and Marketing 16

A MacLeod, M Edney, and MS Izydorczyk

Soybean: Grading and Marketing 25

EG Hammond, LA Johnson, and PA Murphy

Identification of Varieties of Food Grains 29

DM Miskelly and CW Wrigley

Grain Harvest, Storage and Transport 42

Wheat: Harvesting, Transport, and Storage 42
S Grundas and CW Wrigley

Barley: Harvesting, Storage, and Transport 50

RJ Henry

Sorghum: Harvest, Storage, and Transport 54

T Beta, M Chisi, and ES Monyo

Canola: Harvest, Transport, and Storage 62

JJ Mayko

Cereals: Grain Defects 68

SA Barton

The Nature, Causes, and Control of Grain Diseases in the Major Cereal Species 74
RA McIntosh, PM Williamson, and CW Wrigley

Contaminants of Grain 83
J McLean and CW Wrigley
xiv Contents

Chemicals for Grain Production and Protection 99

PC Annis

Handling from Farm to Storage Terminal 105

D Richard-Molard and CW Wrigley

Stored Grain: Invertebrate Pests 110

PC Annis

Postharvest Operations for Quality Preservation of Stored Grain 117

F Fleurat-Lessard

Stored-Grain Pest Management 126

F Fleurat-Lessard

Agronomy of Grain Growing 140

Implication of Climate Changes 140
S Ceccarelli

Sustainable Grain Production and Utilization 144

L Munck

Organic Grain Production and Food Processing 154

P Gelinas and C David

Precision Agriculture 162

EC Leonard

Plants: Diseases and Pests 168

SA Barton

Wheat: Agronomy 176

GM Paulsen, JP Shroyer, and KJ Shroyer

Barley: Agronomy 186

RD Horsley and M Hochhalter

Maize: Agronomy 194

ED Nafziger

Sorghum: Production and Improvement Practices 201

WL Rooney

Canola: Agronomy 207

NJ Mendham and MJ Robertson

Chickpea: Agronomy 216

KHM Siddique and L Krishnamurthy

Lentil: Agronomy 223

T Nleya, A Vandenberg, FL Walley, and D Deneke

Lupin: Agronomy 231

RJ French

Field Pea: Agronomy 240

RJ French

Soybean: Agronomy 251

ED Nafziger

Natural Disease Control in Cereal Grains 257

WK Mousa and MN Raizada

Wheat: Biotrophic Pathogen Resistance 264

RF Park
 Contents xv

Necrotrophic Pathogens of Wheat 273

RP Oliver, K-C Tan, and CS Moffat

Breeding of Grains 279

Wheat Breeding: Exploiting and Fixing Genetic Variation by Selection and Evaluation 279
R DePauw and L OBrien

Barley: Genetics and Breeding 287

A Cuesta-Marcos, JG Kling, AR Belcher, T Filichkin, SP Fisk, R Graebner, L Helgerson, D Herb, B Meints, AS Ross,
PM Hayes, and SE Ulrich

Maize: Breeding 296

EA Lee and LW Kannenberg

Rice: Breeding 304

Qingyao Shu and Dianxing Wu

Canola/Rapeseed: Genetics and Breeding 311

PBE McVetty and RW Duncan

Lentil: Breeding 317

W Erskine, A Sarker, and S Kumar

Lupin: Breeding 325

LC Trugo, E von Baer, and D von Baer

Soybean: Germplasm, Breeding, and Genetics 333

RG Palmer and T Hymowitz

Genetics of Grains 343

Development of Genetically Modified Grains 343
L Privalle

Detection of Genetically Modified Grains 350

RD Shillito

Genomics of Food Grains 360

RJ Henry

Genome Mapping 365

VK Tiwari, JD Faris, B Friebe, and BS Gill

Wheat Genetics 376

RA McIntosh

Wheat Genetics and Genomics 382

E Akhunov

Biotic Stress Resistance Genes in Wheat 388

J Singla and SG Krattinger

Abiotic Stress Genes and Mechanisms in Wheat 393

SJ Roy, NC Collins, and R Munns

Rice: Genetics 398

JS Bao and H Corke

Maize: Genetics 407

EA Lee

Maize: Quality Protein Maize 420

JS Li and SK Vasal

Maize: Other Maize Mutants 425

MH Blanco, H Yangcheng, and J Jane
xvi Contents

Appendix 1. Nutrient-Composition Tables for Grains and for Grain-Based Products 433
SF Schakel, N Van Heel, and J Harnack

Appendix 2. Lists of Standardized Testing Methods for the Analysis of Grain and Grain-Based Foods 450
AR Bridges and CW Wrigley

Appendix 3. Grains, Foods, and Ingredients Suiting Gluten-Free Diets for Celiac Disease 460
JM Jones, V Zevallos, and CW Wrigley

Glossary 467
Index 491
PREFACE

The grain-producing plants are essential to life on earth. They are the primary means by which sunlight, the
primary source of the earths energy, is captured via photosynthesis to turn carbon dioxide and water into
carbohydrates and thus into the wider range of the biochemicals needed by all life forms.
The summary is simple:
The suns energy CO2 H2O
#
Grain-bearing plants
which provide:
Food, Fats, Fuel and Fiber for ourselves
Feed, Forage, Fodder for our animals
Food for Fish Farms

Directly, grains supply over half of humanitys energy and protein.

Indirectly, grains contribute to our food supply via the feeding of grains and grain by-products to animals,
birds and fish.
Grains are increasingly a renewable source of fuels (ethanol and oils).
Grain species, such as cottonseed, contribute fiber for our clothing.

Grains to Feed and Fuel our World

As its title indicates, the Encyclopedia of Food Grains concentrates on the food uses of grains, but details are also
provided about the wider roles of grains. Greater detail about the use of grains for animal feeding is already
available via the on-line Feedipedia encyclopedia (http://www.feedipedia.org/).
Worldwide, however, it is our food that is our obsession first and foremost. We see our health (or
otherwise) to be intimately related to our diet. Therefore nutrition is a major accent of this second edition,
occupying the whole of the second volume, following the first volumes description of the basic aspects of the
world of grains. The third and fourth volumes work their ways back up the value-added grain chain to describe
grain processing and production, thence to breeding and genetics.
The sequence of articles in the first edition (the Encyclopedia of Grain Science, 2004) followed an alphabetical
order. In contrast, the distinct ordering of articles in the second edition assembles fundamental concepts in
Volume 1 The World of Food Grains. Moving into the later volumes, greater complexity and erudition are
evident. Thus, unusual terms such as Transcriptomics, Aeciospore and Allelopathy are more likely to be
found in later volumes than in the first. In any case, reference to the glossary and the index should help in
elucidating such terms.

The First and Second Editions - Editors

This second edition is a considerable expansion on the content of the first edition, with its three print volumes
and 168 articles. Now, some twelve years later, the four print volumes of the Encyclopedia of Food Grains provide

xvii
xviii Preface

216 articles, many newly written for the new edition. Those articles that have been reprinted from the first
edition have been updated. In all cases, authors and reviewers are world renowned experts in their respective
fields.
This second edition has been more than three years in its development. In the early stages, the editors were
Harold Corke, Chuck Walker and Colin Wrigley (the three editors of the first edition). Sadly in 2012, Chuck
Walker passed away, leaving a gap that was difficult to fill.
Chuck was replaced by Koushik Seetharaman, who worked avidly in developing the concepts for this second
edition, including his contributions to a meeting of the editors and Elsevier staff in Sydney, January, 2013.
Sadly, Koushik suffered a heart attack in June, 2014. However, his contributions to the second edition during
this critical development stage were such that Elsevier decided that his name should remain as a co-editor.
Tributes to Professors Walker and Seetharaman are provided below.
In July, 2014, Professor Jon Faubion was appointed as a co-editor. Dr Faubion is the Charles Singleton
Professor of Baking and Cereal Science in the Department of Grain Science, Kansas State University. He is a
more than worthy replacement under such tragic circumstances.

Acknowledgement to All Involved

The production of an encyclopedia involves a wide range of experts with expertise in a diversity of areas of
knowledge. We acknowledge with grateful thanks the groups of contributors:
Members of the Editorial Advisory Board;
Authors who have shared their special knowledge and experience;
Elsevier staff who have provided extensive advice and administrative services, especially Donna de
Weerd-Wilson, Simon Holt and Rachel Gerlis (successive Supervisory Staff) and Gemma Tomalin,
Gemma Taft, Joanne Williams and Mark Harper (successive Content Project Managers).
The Editors
Harold Corke, Jon Faubion, Koushik Seetharaman, and Colin Wrigley
TRIBUTES TO FORMER CO-EDITORS

Professor Emeritus Charles (Chuck) E. Walker

Chuck Walker was one of the three editors of the first edition of Elseviers Grains
Encyclopedia (the Encyclopedia of Grain Science, 2004). He was again appointed
as an editor of this second edition. He participated in the planning stages for the
second edition until his untimely death on 26 April 2012. At the time of his
death, Chuck was a Professor Emeritus at the Department of Grain Science and
Industry at Kansas State University, Manhattan, KS, USA.
Charles Eugene Walker was born on 17 December 1936, in Winterset, Mad-
ison county, IA, USA, during the great depression. He was the eldest child of
Albert Cecil and Bessie Irene Headley Walker, poor farmers in Winterset, Mad-
ison county, IA, USA. He walked more than a mile to attend rural one-room
grammar schools in Warren county and graduated from high school in New
Virginia, IA, USA.
Better known as Chuck in later life, Dr Walker received a BS degree in
Chemical Engineering from Iowa State University, Ames, IA, USA, in May 1959. At a mixer party in the fall
of 1957, he met Shannon Rose Philp, a pretty freshman student from Loup city, NE, USA. This was just after a
summer spent working alone on a fire-lookout tower for the US Forest Service, in the wilds of Clearwater
National Forest, in Northern Idaho. They were married on 1 June 1958 and she became his wife of 54 years. She
later rewarded him with two sons, Alan Eugene and Tomas Charles, and they adopted a daughter, Bekianne.
Upon graduation, Chuck worked in flour milling research at the General Mills Central Research Laboratories
in the Minneapolis area from 1959 to 1962. He then accepted a fellowship to graduate school at North Dakota
State University in Fargo from 1962 to 1965, majoring in Cereal Chemistry. Chuck received his PhD in
Chemistry in the summer of 1966. For his PhD dissertation, he developed a micro brewery and studied barley
proteins and their influence on beer quality a rather ironic choice of topics, considering he remained a strict
teetotaler all his life.
While finishing his thesis, he joined the faculty of the North Dakota State College, Valley city, ND, USA
(19651974), teaching various physical science and chemistry courses to undergraduate students. Feeling the
pull of industry, he moved south to work for the Fairmont Foods Central Research Laboratory in Omaha, NE,
USA (19741980), where he did research, product development, and technical assistance on baking, pizza,
snack, and dairy products, eventually reaching the position of Associate Director of Research.
Dismemberment and sale of this Fortune-500 company provided him an opportunity to return to academia
at the University of Nebraska, Lincoln, NE, USA (19801987), where he developed a Cereal Technology
Program within the Department of Food Technology. He developed and taught various food science and
technology courses to both undergraduate and graduate students, and developed a vigorous research program.
He became Interim Department Head and Food Processing Center Director.
In December 1987, Chuck joined the Department of Grain Science and Industry at Kansas State University as
the Bakers National Educational Foundations Baking Science Professor. There he maintained a busy program

xix
xx Tributes to Former Co-Editors

in research, teaching, graduate student advising, publishing, traveling and consulting, specializing in baking
technology.
During his career, Chuck published more than 100 technical papers, and advised about 50 MS and PhD
graduate students. He was heavily involved in applied rheology (mixing and flow characteristics) of flour
products. In recognition of his commitment to international graduate students, Mr Roland Temme, TMCO,
Lincoln, NE, USA, established an international graduate student fellowship in his honor.
Chuck entered 50% phased-in retirement at age 65 in 2002 and retired in 2005, only to be hired back 1/
10-time afterward to assist with seminars and the departments centennial recognition in 2010. He continued
to support the departments activities on a volunteer basis.
Chucks research and consulting travels took him to more than a dozen countries, including many extended
trips to Australia, eventually earning permanent resident status in Australia a country which had fascinated
him since childhood. Most of his time in Australia was spent with the Bread Research Institute in Sydney. He
also made several extended trips to China, serving as guest lecturer at ShanDong Agricultural University, TaiAn,
and at Henan University of Technology, Zhengzhou.
Chuck always commented that the best legacy a person could leave is the knowledge he has shared. A
significant way in which his knowledge has been shared is represented by his contributions to the first and
second editions of Elseviers Encyclopedia of Food Grains.

Colin Wrigley on behalf of the Editors.

Professor Koushik Seetharaman

In August 2012, Koushik Seetharaman was appointed as the third editor of this
second edition of Elseviers Grains Encyclopedia, replacing Professor Walker.
Tragically, Koushiks many contributions to the Encyclopedia of Food Grains were
cut short by his untimely death at 48 years in June 2014.
Koushik received his BS at the Gujarat Agricultural University, Anand, Guja-
rat, India. His MS training was at Cornell University, Ithaca, NY, USA, and his
PhD research was carried out at Texas A&M University, College Station, TX, USA.
At the time of Koushiks appointment as encyclopedia editor (August 2012),
he was Associate Professor and Cereals Chair in the Department of Food Science
at the University of Guelph, Canada. Research in his laboratory spanned the
cereal-grain value chain, focusing on grain quality, processing and ingredient
interactions, and on consumer acceptability and health.
His research on grain quality at Guelph involved collaboration with cereal-breeding companies, grain
farmers of Ontario and local processors, by analyzing Ontario wheats for functionality and nutritional
attributes. Special accents were on antioxidant activity, and on fiber and phenolics contents. Interactions
with breeders and processor related to the chemistry, sensory properties and consumer acceptance of whole
grain products, with specific focus on red versus white wheats in different product matrices.
Koushik had a special interest in the behavior of starch in water-poor systems, such as dough and baked
products. His research group had demonstrated that the constituent polymers of wheat starch are a series of
biopolymers ranging from linear to branched, depending on several factors including genetics and growth
environment. Moreover, the proportion of these polymers further defines their functional properties.
Research funding had been provided by the Canadian Agricultural Adaptation Council with contributions
from the Ontario Cereal Industry Research Council, Kellogg, Kraft, Kraft Canada Mill, Dow AgroSciences, C&M
Seeds, Grain Farmers of Ontario and Brabender GmbH.
Several months before his death, Koushik had been appointed as an Associate Professor to the Department
of Food Science and Nutrition at the University of Minnesota, St Paul, MN, USA, there taking the position of
General Mills Cereal Chair. In that capacity, Koushik had again initiated collaborations with breeders and
geneticists, millers and processors, with a focus on what product quality means for the consumer. His research
focused on interactions of grain biopolymers starch and gluten in particular and the impact on processing,
product attributes and consumer health.
 Tributes to Former Co-Editors xxi

His group included three postdoctoral research associates, five doctoral students and one MS student. They
were collaborating extensively with research groups around the world including University of Milan, Italy;
University of Copenhagen, Denmark; Lille University, France; ONIRIS, France; Tamilnadu Agricultural Uni-
versity, India; Iowa State University, USA; African Rice Research Institute, Benin; University of Guelph, Canada;
and Memorial University, Newfoundland, Canada.
Koushiks research background and his collaborations placed him well to contribute to the preparation of
the grains encyclopedia, both by his our writing and via his contacts to engage world-authority contributors.
The grain-science community world-wide was shocked by Koushiks sudden passing. Koushik left a wife
Debra Freedman and an eight-year-old son, Samuel.
Koushiks contribution to the encyclopedia was such that Elsevier decided that his name should remain as a
co-editor even though he did not live to see the end result of his contributions.

Colin Wrigley on behalf of the Editors.

This page intentionally left blank
EDITORS BIOGRAPHY

Colin W. Wrigleys 55 years in grain-science research have earned him international recognition in
the form of several international and Australian research awards. His work is described in about
600 research publications, including several patents, a series of eight books on Australian cereal
varieties, and many edited books. He was Editor-in-Chief of the first edition of the Encyclopedia of
Grain Science (2004).
His research interests have centered on the characterization of cereal-grain proteins in relation to
processing quality. This has involved developing new methods of protein fractionation, including
gel isoelectric focusing and its 2D combination with gel electrophoresis, leading into proteomic
mapping. Other diagnostic methods developed relate to the evaluation of grain quality in wheat
and barley, such as better methods for variety identification and for characterizing quality in starch
and sprouted grain (as co-patentee of the Rapid ViscoAnalyser). Research involvement has also
included elucidation of grain-quality variation due to environmental factors (heat stress, fertilizer
use, CO2 levels and storage conditions).
In 2009, Wrigley was appointed a Member of the Order of Australia (AM) for service to primary
industry, particularly to grain science as a researcher, and to the development of methods for
improving wheat quality.
He is currently an Honorary Professor at the University of Queensland, Brisbane, Australia.

Harold Corke is a professor in the Food and Nutritional Sciences program at The University of
Hong Kong, and a Chutian Scholar Distinguished Foreign Professor in the Glyn O. Phillips
Hydrocolloid Research Center at HUT, Hubei University of Technology, Wuhan, China. In more
than 22 years in Hong Kong, he has had responsibility for teaching a diverse array of food science
courses, including the major courses Grain Production and Utilization, Food Safety and Quality
Management, Food Product Development, and general education courses Feeding the World, and
Food: Technology, Trade and Culture. He is author or co-author of 180 refereed journal articles,
and his 22 PhD graduates have gone on to successful careers in academia and industry around the
world. He is on the editorial boards of Journal of Food Composition and Analysis, LWT Food Science and
Technology and Journal of Cereal Science, and with Colin Wrigley and Chuck Walker was one of the
editors of the first edition of Encyclopedia of Grain Science, published in 2004. Corke is active in
consulting in grain processing and food safety in Asia and Eastern Europe.

Jon Faubion is the Charles Singleton Professor of Baking and Cereal Science in the Department of
Grain Science, Kansas State University. Over his 35 year career, he has had the sole or shared
responsibility for teaching nine different cereal or food science courses at Texas A&M University,
The University of Minnesota and Kansas State. In 2011, he received the Excellence in Teaching
Award from the American Association of Cereal Chemists International. He is a senior editor of
Cereal Chemistry, and a member of the Board of Directors of the American Association of Cereal
Chemists International and an executive editor of Journal of the Science of Food and Agriculture. Prior
to rejoining the department faculty in 2005, he directed the Applied Technology and Sensory
Science Groups for the research and development arm of The Schwan Food Company.

xxiii
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EDITORIAL ADVISORY BOARD

Jinsong Bao Bob French

Zhejiang University, Hangzhou, PR China
Frank Bekes
Director R&D FBFD Pty Ltd, Beecroft, NSW, Australia
Trust Beta
University of Manitoba, Winnipeg, MB, Canada
Stanley Cauvain
BakeTran, Witney, United Kingdom; Curtin University,
Perth, WA, Australia
Robert Cracknell
Crackers Consulting, Mount Eliza, VIC, Australia
Yapeng Fang
Hubei University of Technology, Wuhan, PR China
Jon Faubion
Kansas State University, Manhattan, KS, USA
Department of Agriculture and Food, Western Australia,
Merredin, WA, Australia
Robert Graybosch
Agricultural Research Service, USDA, Lincoln, NE, USA
Robert Henry
University of Queensland, Brisbane, QLD, Australia
Barry V. McCleary
Megazyme International Ireland, Bray, Wicklow, Ireland
Lindsay OBrien
University of Sydney, Narrabri, NSW, Australia
Paul Scott
USDA-ARS Corn Insects and Crop Genetics Research
Unit, Ames, IA, USA
John Reginald Nuttall Taylor
University of Pretoria, Pretoria, South Africa

xxv
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CONTRIBUTORS

E Akhunov BJ Blaney
Kansas State University, Manhattan, KS, USA The University of Queensland, St Lucia, Australia
PC Annis JE Bock
CSIRO Entomology, Canberra, ACT, Australia University of Guelph, Guelph, ON, Canada
GA Annor F Bonomi
University of Ghana, Accra, Ghana University of Milan, Milan, Italy
CA Atkins AR Bridges
The University of Western Australia, Crawley, WA, AACC International, St Paul, MN, USA
Australia
H Brinch-Pedersen
W Atwell Aarhus University, Slagelse, Denmark
Bill Atwell Consulting, Champlin, MN, USA
G Bultosa
M Baga Haramaya University, Dire Dawa, Ethiopia; Botswana
University of Saskatchewan, Saskatoon, SK, Canada College of Agriculture, Gaborone, Botswana
JS Bao W Bushuk
Zhejiang University, Hangzhou, PR China University of Manitoba, Winnipeg, MB, Canada
VJ Barthet YZ Cai
Canadian Grain Commission, Winnipeg, MB, Canada China National Seed Group, Beijing, PR China;
SA Barton Huazhong Agricultural University, Wuhan, PR China;
Oxford Agricultural Trials Ltd, Oxford, UK The University of Hong Kong, Hong Kong, PR China;
Hubei University of Technology, Wuhan, PR China
IL Batey
Sunset Cereal Services, Meadowbank, NSW, Australia EF Caldwell
AACC International, Roseville, MN, USA
F Bekes
FBFD PTY LTD, Beecroft, NSW, Australia KJ Carpenter
University of California, Berkeley, CA, USA
AR Belcher
Oregon State University, Corvallis, OR, USA JC Catullo
INTA, Buenos Aires, Argentina
JN BeMiller
Purdue University, West Lafayette, IN, USA SP Cauvain
BakeTran, Witney, OX, UK
T Beta
University of Manitoba, Winnipeg, Manitoba, Canada S Ceccarelli
Via delle Begonie 2, Ascoli Piceno, Italy
JL Black
John L Black Consulting, Warrimoo, NSW, Australia GK Chandi
University of Guelph, Guelph, ON, Canada
MH Blanco
(Retired) Plant Breeder and Geneticist, Encinitas, N Chaudhary
CA, USA G. J. University of Science and Technology, Hisar, India

xxvii
xxviii Contributors

RN Chibbar JE Dexter
University of Saskatchewan, Saskatoon, SK, Canada Canadian Grain Commission, Winnipeg, MB, Canada
M Chisi B Dobraszczyk
Golden Valley Research Station, Chisamba, Zambia Reading Science Centre, Reading, UK
S Cloutier D Domingues
Agriculture and Agri-Food Canada, Ottawa, ON, General Mills Inc., Minneapolis, MN, USA
Canada
C Dowd
LS Collado General Mills Inc., Minneapolis, MN, USA
The University of the Philippines Los Banos, Laguna,
LM Duizer
Philippines
University of Guelph, Guelph, ON, Canada
NC Collins
RW Duncan
School of Agriculture Food and Wine, The University of
University of Manitoba, Winnipeg, MB, Canada
Adelaide, Glen Osmond, SA, Australia
KG Duodu
H Corke
University of Pretoria, Pretoria, South Africa
The University of Hong Kong, Hong Kong, Pokfulam,
PR China; Hubei University of Technology, Wuhan, SR Eckhoff
PR China (Retired) University of Illinois, Urbana, IL, USA
RL Cracknell M Edney
Crackers Consulting, Mount Eliza, VIC, Australia Canadian Grain Commission, Winnipeg, MB, Canada
GB Crosbie M Emery
Crosbie Grain Quality Consulting, East Fremantle, WA, Iowa State University, Ames, IA, USA
Australia
W Erskine
JS Croser University of Western Australia, Crawley, WA, Australia
University of Western Australia, Crawley, WA, Australia
WD Eustace
A Cruickshank (Retired) Kansas State University, Manhattan, KS, USA
DAFF Queensland, Warwick, QLD, Australia
A Evans
A Cuesta-Marcos Tate and Lyle, Hoffman Estates, IL, USA
Oregon State University, Corvallis, OR, USA
E Evans
MG DEgidio The Tassie Beer Dr, Lindisfarne, TAS, Australia
Unita di ricerca per la Valorizzazione Qualitativa dei
JD Faris
Cereali, Rome, Italy
USDA-ARS Cereal Crops Research Unit, Fargo, ND,
D Dan Ramdath USA
Agriculture and Agri-Food Canada, Guelph, ON, Canada
J Faubion
P Dangi Kansas State University, Manhattan, KS, USA
G. J. University of Science and Technology, Hisar, India
AM Fehily
JK Daun Tinuviel Software, Warrington, UK
(Deceased)
T Filichkin
C David Oregon State University, Corvallis, OR, USA
ISARA Lyon, Lyon, France
GB Fincher
L Day Australian Research Council Centre of Excellence in Plant
AgResearch Ltd., Palmerston North, New Zealand Cell Walls, University of Adelaide, Glen Osmond, SA,
Australia
D Deneke
South Dakota State University, Brookings, SD, USA SP Fisk
Oregon State University, Corvallis, OR, USA
R DePauw
Agriculture and AgriFood Canada, Swift Current, SK, MT Fletcher
Canada The University of Queensland, St Lucia, Australia
 Contributors xxix

RJ Fletcher A Gunaratne
University of Queensland, Gatton, QLD, Australia The University of Hong Kong, Hong Kong, PR China
F Fleurat-Lessard YN Guragain
INRA, Mycology and Food Safety Research Unit, Kansas State University, Manhattan, KS, USA
Villenave dOrnon, France
RCE Guy
JA Fox Campden and Chorleywood Food Research Association,
Kansas State University, Manhattan, KS, USA Chipping Campden, UK
RJ French C Hall III
The University of Western Australia, Crawley, WA, North Dakota State University, Fargo, ND, USA
Australia
LM Hall
B Friebe University of Alberta, Edmonton, AB, Canada
Kansas State University, Manhattan, KS, USA
EG Hammond
BJ Furman Iowa State University, Ames, IA, USA
San Jose State University, San Jose, CA, USA
R Haraszi
M Gangola Campden BRI, Gloucestershire, UK
University of Saskatchewan, Saskatoon, SK, Canada
J Harnack
P Gelinas University of Minnesota, Minneapolis, MN, USA
Food Research and Development Centre, Agriculture and
D Hauge
Agri-Food Canada, Saint-Hyacinthe, QC, Canada
Grains for Health Foundation, St. Louis Park, MN, USA
SM Ghazani
AMR Hayes
University of Guelph, Guelph, ON, Canada
St. Catherine University, Lafayette, MN, USA
MC Gianibelli
PM Hayes
(Retired)
Oregon State University, Corvallis, OR, USA
A Gianinetti
L Helgerson
Consiglio per la ricerca in agricoltura e lanalisi
Oregon State University, Corvallis, OR, USA
delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Italy RJ Henry
University of Queensland, Brisbane, QLD, Australia
BS Gill
Kansas State University, Manhattan, KS, USA D Herb
Oregon State University, Corvallis, OR, USA
JS Godber
LSU Agricultural Center, Baton Rouge, LA, USA I Herencia
Universidad Politecnica de Madrid, Madrid, Spain
A Goldstein
University of Minnesota, St. Paul, MN, USA E Hernandez
Texas A&M University, College Station, TX, USA
R Graebner
Oregon State University, Corvallis, OR, USA KB Hobson
New South Wales Department of Primary Industries,
S Graham-Acquaah
Tamworth, NSW, Australia
Africa Rice Center, Cotonou, Republic of Benin
M Hochhalter
RA Graybosch
North Dakota State University, Fargo, ND, USA
USDA-ARS, University of Nebraska Lincoln, Lincoln,
NE, USA RD Horsley
North Dakota State University, Fargo, ND, USA
S Grundas
Bohdan Dobrzanski Institute of Agrophysics of the Polish W Huang
Academy of Sciences, Lublin, Poland Jiangsu Academy of Agricultural Sciences, Nanjing,
PR China; Jiangnan University, Wuxi, Jiangsu, PR China
TJ Gulya
USDA-ARS, Northern Crop Science Laboratory, Fargo, Y Huang
ND, USA China Agricultural University, Beijing, PR China
xxx Contributors

V Humphrey-Taylor P Koehler
(Retired) New Zealand Institute for Crop Food Research Deutsche Forschungsanstalt fur Lebensmittelchemie,
Limited, Christchurch, New Zealand Leibniz Institut, Freising, Germany
T Hymowitz R Korczak
University of Illinois at Champaign-Urbana, Urbana, University of Minnesota, St. Paul, MN, USA
IL, USA
SG Krattinger
S Iametti University of Zurich, Zurich, Switzerland
University of Milan, Milan, Italy
L Krishnamurthy
C Isaak International Crops Research Institute for the Semi-Arid
University of Manitoba, Winnipeg, MB, Canada Tropics, Patancheru, Andhra Pradesh, India
MS Izydorczyk S Kumar
Canadian Grain Commission, Winnipeg, MB, Canada International Center for Agricultural Research in the Dry
Areas (ICARDA), Rabat, Morocco
P Jacques
Tufts University, Boston, MA, USA OR Larroque
CSIRO Agriculture Flagship, Canberra, Australia
S Jaiswal
University of Saskatchewan, Saskatoon, SK, Canada JW Lawton
J Jane (Retired) ARS-USDA
Iowa State University, Ames, IA, USA EA Lee
LA Johnson University of Guelph, Guelph, ON, Canada
Iowa State University, Ames, IA, USA EC Leonard
JM Jones AgriKnowHow, Maitland, SA, Australia
St. Catherine University, Arden Hills, MN, USA JS Li
A Juhasz China Agricultural University, Beijing, PR China
Centre for Agricultural Research of the Hungarian WD Li
Academy of Sciences, Martonvasar, Hungary Shanxi University, Taiyuan, PR China
BO Juliano Z Li
Philippine Rice Research Institute Los Banos, Laguna, CSIRO Food Futures National Research Flagship,
Philippines Canberra, ACT, Australia; CSIRO Plant Industry,
RS Kadan Canberra, ACT, Australia
(Deceased)
R Lindberg
GP Kadkol Minneapolis Heart Institute Foundation, Minneapolis,
NSW Department of Primary Industries, Calala, NSW, MN, USA
Australia
MG Lindhauer
LW Kannenberg Max Rubner-Institut (MRI), Detmold, Germany
University of Guelph, Guelph, ON, Canada
K Liu
TN Khan Agricultural Research Service, US Department of
University of Western Australia, Crawley, WA, Australia Agriculture, Aberdeen, ID, USA

BS Khatkar Q Liu
G. J. University of Science and Technology, Hisar, India Agriculture and Agri-Food Canada, Guelph, ON, Canada

JG Kling S Lu
Oregon State University, Corvallis, OR, USA National Chung Hsing University, Taichung, Taiwan
EJ Knights OM Lukow
New South Wales Department of Primary Industries, Agriculture and Agri-Food Canada, Winnipeg, MB,
Tamworth, NSW, Australia Canada
 Contributors xxxi

BL Ma NJ Mendham
Agriculture and Agri-Food Canada, Ottawa, ON, University of Tasmania, Hobart, TAS, Australia
Canada
R Menon
C-Y Ma The General Mills Bell Institute of Health and Nutrition,
The University of Hong Kong, Hong Kong, PR China Minneapolis, MN, USA
A MacLeod TE Michaels
Canadian Grain Commission, Winnipeg, MB, Canada University of Minnesota, St. Paul, MN, USA
L MacLeod DM Miskelly
Dairy Innovation Australia, Werribee, VIC, Australia Westcott Consultants P/L, Goulburn, NSW, Australia
CK Madsen
CS Moffat
Aarhus University, Slagelse, Denmark
Curtin University, Bentley, WA, Australia
RJ Mailer
ES Monyo
Australian Oils Research, Lambton, NSW, Australia
ICRISAT, Nairobi, Kenya
JT Manful
Africa Rice Center, Cotonou, Republic of Benin TR Moore
AIB International, Manhattan, KS, USA
AG Marangoni
University of Guelph, Guelph, ON, Canada MK Morell
International Rice Research Institute, Los Banos,
DJ Mares Philippines
University of Adelaide, Glen Osmond, SA, Australia
CF Morris
L Marquart USDA-ARS Western Wheat Quality Laboratory,
University of Minnesota, St. Paul, MN, USA; Grains for Pullman, WA, USA
Health Foundation, St. Louis Park, MN, USA
LA Morrison
A Marti
Oregon State University, Corvallis, OR, USA
Universita degli Studi di Milano, Milan, Italy
B Maschoff WK Mousa
Grains for Health Foundation, St. Louis Park, MN, USA University of Guelph, Guelph, ON, Canada; Mansoura
University, Mansoura, Egypt
A Mathiowetz
University of Minnesota, St. Paul, MN, USA; Rochester, K Mrva
MN, USA University of Adelaide, Glen Osmond, SA, Australia

JJ Mayko L Munck
Canola Council of Canada, Mundare, AB, Canada University of Copenhagen, Frederiksberg C, Denmark
CM McDonough R Munns
Texas A&M University, College Station, TX, USA CSIRO Agriculture, Canberra, ACT, Australia; The
University of Western Australia, Crawley, WA, Australia
RA McIntosh
The University of Sydney, Cobbitty, NSW, Australia PA Murphy
Iowa State University, Ames, IA, USA
JD McKeehen
Cereal Partners Worldwide, Minneapolis, MN, USA ED Nafziger
J McLean University of Illinois, Urbana, IL, USA
The University of Newcastle, Callaghan, NSW, Australia SC Nelson
PBE McVetty AACC International (AACCI), St. Paul, MN, USA
University of Manitoba, Winnipeg, MB, Canada T Nleya
B Meints South Dakota State University, Brookings, SD, USA
Oregon State University, Corvallis, OR, USA
L OBrien
A Meldrum The University of Sydney, Narrabri, NSW, Australia;
Pulse Australia, Perth, WA, Australia Solheimar Pty Ltd, Narrabri, NSW, Australia
xxxii Contributors

DK OToole I Rajcan
City University of Hong Kong, Hong Kong, PR China University of Guelph, Guelph, ON, Canada
EA Oelke L Ramsden
University of Minnesota, St. Paul, MN, USA University of Hong Kong, Hong Kong, PR China
RP Oliver KD Rausch
Curtin University, Bentley, WA, Australia University of Illinois at UrbanaChampaign, Urbana, IL,
USA
DE Ortiz
W. K. Kellogg Institute, Battle Creek, MI, USA A Regina
MA Pagani CSIRO Food Futures National Research Flagship,
Universita degli Studi di Milano, Milan, Italy Canberra, ACT, Australia; CSIRO Plant Industry,
Canberra, ACT, Australia
TS Palav
Rich Products Corporation, Buffalo, NY, USA MN Riaz
Texas A&M University, College Station, TX, USA
L Pallas
Yanco Agricultural Institute, Yanco, NSW, Australia D Richard-Molard
(Retired) INRA, Nantes, France
GH Palmer
Heriot-Watt University, Edinburgh, UK F Rizza
Consiglio per la ricerca in agricoltura e lanalisi
RG Palmer delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Iowa State University, Ames, IA, USA Italy
RF Park MJ Robertson
The University of Sydney, Narellan, NSW, Australia CSIRO Sustainable Ecosystems, St. Lucia, QLD,
GM Paulsen Australia
Kansas State University, Manhattan, KS, USA LW Rooney
LC Paulson Texas A&M University, College Station, TX, USA
The Bama Companies, Tulsa, OK, USA WL Rooney
DS Petterson Texas A&M University, College Station, TX, USA
Department of Agriculture, Nedlands, WA, Australia CM Rosell
RA Porter Institute of Agrochemistry and Food Technology (IATA-
University of Minnesota-NCROC, Grand Rapids, CSIC), Paterna, Valencia, Spain
MN, USA AS Ross
L Privalle Oregon State University, Corvallis, OR, USA
Bayer CropScience, Research Triangle Park, Durham, SJ Roy
NC, USA School of Agriculture Food and Wine, The University of
KV Probst Adelaide, Glen Osmond, SA, Australia
Kansas State University, Manhattan, KS, USA; Grain HD Sapirstein
Processing Corporation, Muscatine, IA, USA University of Manitoba, Winnipeg, MB, Canada
KJ Quail AK Sarkar
Grain Growers Limited, North Ryde, NSW, Australia Canadian International Grains Institute, Winnipeg, MB,
RCN Rachaputi Canada
University of Queensland, Kingaroy, QLD, Australia
A Sarker
H Rahman International Center for Agricultural Research in the Dry
University of Alberta, Edmonton, AB, Canada Areas (ICARDA), New Delhi, India
S Rahman SK Sathe
Monash University Malaysia, Bandar Sunway, Malaysia Florida State University, Tallahassee, FL, USA
MN Raizada SF Schakel
University of Guelph, Guelph, ON, Canada University of Minnesota, Minneapolis, MN, USA
 Contributors xxxiii

KA Scherf VA Solah
Deutsche Forschungsanstalt fur Lebensmittelchemie, Curtin University, Perth, WA, Australia
Leibniz Institut, Freising, Germany
AM Stanca
MP Scott University of Modena and Reggio Emilia, Reggio Emilia,
Iowa State University, Ames, IA, USA Italy
K Seetharaman FL Stoddard
(Deceased) University of Helsinki, Helsinki, Finland
GJ Seiler K-C Tan
USDA-ARS, Northern Crop Science Laboratory, Fargo, Curtin University, Bentley, WA, Australia
ND, USA
AS Tatham
SO Serna-Saldivar Cardiff Metropolitan University, Cardiff, UK
Centro de Biotecnologa FEMSA, Escuela de Ingeniera y
JRN Taylor
Ciencias, Monterrey, Mexico
University of Pretoria, Hatfield, South Africa
S Shea Miller
V Terzi
Eastern Cereals and Oilseeds Research Centre,
Consiglio per la ricerca in agricoltura e lanalisi
Agriculture and Agri-Food Canada, Ottawa, ON,
delleconomia agraria, CRA-GPG, Fiorenzuola dArda,
Canada
Italy
DB Sheats
K Tiefenbacher
St. Catherine University, St. Paul, MN, USA
(Retired)
PR Shewry
VK Tiwari
Rothamsted Research, Harpenden, UK; University of
Kansas State University, Manhattan, KS, USA
Reading, Reading, UK
SM Tosh
RD Shillito
Agriculture and Agri-Food Canada, Guelph, ON, Canada
Bayer CropScience, Durham, NC, USA
P Tosi
JP Shroyer
University of Reading, Reading, UK
Kansas State University, Manhattan, KS, USA
LC Trugo
KJ Shroyer
Federal University of Rio de Janeiro, Rio de Janeiro, Brazil
Kansas State University, Manhattan, KS, USA
SE Ulrich
Qingyao Shu
Washington State University, Pullman, WA, USA
Zhejiang University, Hangzhou, PR China
EH Unger
KHM Siddique
Eagle Rock Solutions, Inc., Idaho Falls, ID, USA
University of Western Australia, Crawley, WA, Australia
PV Vadlani
J Singla
Kansas State University, Manhattan, KS, USA
University of Zurich, Zurich, Switzerland
SA Valencia-Chamorro
M Sissons
Escuela Politecnica Nacional, Quito, Ecuador
NSW Department of Primary Industries, Calala, NSW,
Australia; Tamworth Agricultural Institute, Tamworth, V Vamadevan
NSW, Australia University of Minnesota, St. Paul, MN, USA; Agriculture
and Agri-Food Canada, Guelph, ON, Canada
JH Skerritt
University of Queensland, St Lucia, QLD, Australia N Van Heel
University of Minnesota, Minneapolis, MN, USA
DJ Skylas
Grain Growers Limited, North Ryde, NSW, A Vandenberg
Australia University of Saskatchewan, Saskatoon, SK, Canada
DL Smith SK Vasal
McGill University, Ste-Anne-de-Bellevue, QC, Canada BISA, Punjab, India
xxxiv Contributors

R Velicogna BJB Wood

Archer Daniels Midland, Montreal, QC, Canada Strathclyde University, Glasgow, UK
D von Baer G Wright
Seeds Baer, Temuco, Chile Peanut Company of Australia, Kingaroy, QLD, Australia

E von Baer CW Wrigley

CAMPEX Semillas Baer, Temuco, Chile
CE Walker
(Deceased)
SB Walker
University of Guelph, Guelph, ON, Canada
FL Walley
University of Saskatchewan, Saskatoon, SK, Canada
D Wang
Huazhong Agricultural University, Wuhan, PR China
T Wang
Iowa State University, Ames, IA, USA
RD Waniska
Texas A&M University, College Station, TX, USA
LL Ward
Kansas State University, Manhattan, KS, USA
H Wieser
Deutsche Forschungsanstalt fur Lebensmittelchemie,
Leibniz Institut, Freising, Germany
RM Williams
CBH Group, West Perth, WA, Australia
PM Williamson
(Retired) Leslie Research Centre, Toowoomba, QLD,
Australia; CBH Group, West Perth, WA, Australia
Honorary Professor, QAAFI; The University of
Queensland, Brisbane, QLD, Australia
Dianxing Wu
Zhejiang University, Hangzhou, PR China
HX Wu
Wuhan Chortle Biotechnology Company Limited,
Wuhan, PR China
J Wyatt
NIAB, Cambridge, UK
Y Yan
Huazhong University of Science and Technology, Wuhan,
PR China
H Yangcheng
Iowa State University, Ames, IA, USA
V Zevallos
Johannes Gutenberg Mainz, Mainz, Germany; University
Medical Center, Mainz, Germany
XM Zhou
McGill University, Ste-Anne-de-Bellevue, QC, Canada
PK Zwer
South Australian Research and Development Institute
(SARDI), Urrbrae, SA, Australia
S Zydenbos
(Retired) New Zealand Institute for Crop Food Research
Limited, Christchurch, New Zealand
 NUTRITION AND FOOD GRAINS
Food Grains and Well-Being
Contents
Functional Foods: Overview
Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics
Nutrition: Soy-Based Foods
Functional Foods: Overview
G Bultosa, Botswana College of Agriculture, Gaborone, Botswana; Haramaya University, Dire Dawa, Ethiopia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Functional food concepts/definitions.
Bioactive compounds.
Chronic diseases and functional foods.
Gluten-free foods for celiac patients.
Learning Objectives
To achieve understanding on the concepts/definitions of
functional foods.
To achieve understanding of bioactive compounds used in
functional foods, grain dietary sources, and potential effects
on health.
To impart processing principles on grain-based functional
foods.
Introduction
Humankind has progressed from a hunter-gather approach to
food consumption to sedentary agriculture, progressing through
the industrial revolution to our current knowledge-based society
with advances in food processing. There is now enormous evi-
dence about the limitations of consuming specific food
component(s) on human health. The twenty-first century of
humankind is marked by lifestyle changes resulting in the con-
sumption of diets high in calories, fat, cholesterol, and sodium
and inadequate intakes of calcium, iron, zinc, and dietary fibers
and an increased resistance of pathogens to drugs. These factors,
coupled with less physical exercise, aging populations, and
more leisure time, have exposed much of the worlds population
to cancers and various metabolic syndrome-related diseases,
osteoporosis, dementia, etc. Some genetically predisposed
individuals are also affected by diet-caused allergens.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00071-1
Consumption of an adequately balanced diet is a means of
body structure formation, energy generation, and health. More
than 2500 years ago, Let foods be our medicine and medicine
be our foods was stated by Hippocrates. This shows that
consumption of diets with health-promoting effects is not
new. But evidence on the relationship between dietary chem-
ical component(s) and health is on evolutionary development
as technology and human comprehension advance. Within
such evolution, the concept of functional foods was started
in the 1980s as Foods for Specified Health Use (FOSHU)
in Japan. Foods, when consumed as a regular diet that supplies
one or more bioactive components beyond basic nutrients and
offer health-promoting effects, are today branded as functional
foods. Functional foods are not prescribed drugs, dietary sup-
plements, medical foods of therapeutic effects, traditional
medicines, or nutraceuticals. Functional foods are distinct
from macronutrient and micronutrient supplements targeted
to achieve balanced diets and to treat nutrient deficiency syn-
dromes. Functional foods thus comprise whole foods, forti-
fied, enriched, or enhanced foods bearing bioactive
compounds beyond macro- and micronutrients. Sources of
bioactive ingredients include whole grains, flaxseed, sesame
seed, psyllium seed husks, legumes (soybean and fenugreek),
fruits, vegetables, herbs, spices, and fermented foods. The
nature of bioactive compounds derived from such sources is
variable even though similarities also exist. Those recognized
bioactive compounds include phenolics; carotenoids; dietary
fibers; b-glucans and inulin-type fructans; o-3 fatty acids; pro-
biotics, prebiotics, and synbiotics; phytoestrogens; soy
proteins; plant stanols and sterols; isothiocyanates; polyols;
and some minerals and vitamins. In some respects, there is
an overlap with food bioactives recognized to be functional
ingredients and micronutrients such as in the case of vitamins
and minerals. Most bioactive compounds present in functional
foods are not necessarily essential for life but are recognized
contributors toward good health.
1
2 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview
The World Health Organization (WHO) recommends to the
public to move toward healthful diets. The functional-food
expansion across the globe is enormous with the current market
size estimated from 7 to 190 billion USD. The ever-increasing
health-care costs and public awareness on roles of diet on health
are contributing factors for such expansion. Functional-food
development involves evaluation of the roles, safety, and con-
sumer acceptance of bioactive compounds. For foods that have
no history of consumption as regular diets (novel foods), the
evaluation and regulatory requirements are rigorous and are
required to be substantiated by scientific evidence for their
efficacy and safety. As a result, even though a number of novel
foods were developed in the past, their wider expansion was
limited.
Various grain-based functional foods are today on the mar-
ket in the form of whole grain snacks, baked foods, extruded
products, ready-to-eat breakfast cereals, fermented cereals, bev-
erages, and gruels. Other products are: dietary fibers, inulin-
type fructans, b-glucans, carotenoids, o-3 fatty acids, plant
sterols, and/or stanols-fortified products. There are also indig-
enous foods that are a heritage of a given community and their
record as safe diet is lost in antiquity and yet most of them
fulfill what is expected of functional foods.
In this article, functional food definitions, distinction from
related concepts, recognized bioactive ingredients, dietary
sources, health benefits, and processing principles are
described giving emphasis on grains. Read also functional
foods: dietary fibers, prebiotics, probiotics, and synbiotics as
these are not covered here.
Functional Food: Concepts and Definitions
So far, there is no global consensus on the definition of func-
tional foods. The Codex Alimentarius Commission only issued
guidelines (CAC/GL 231997) on nutrition and health claims
to be used through consultation with country policy. However,
food regulatory agencies and various professional societies in
different countries have described closely related concepts/
working definitions. Definitions offered from various sources
for functional foods, features, and requirements on what con-
stitute to be nutrition, health, and structure/function claims
are listed under the Further Reading section.
According to the ADA, all foods are functional at some
physiological level because they provide nutrients or other
substances that furnish energy, sustain growth, or maintain/
repair vital processes. However, functional foods are recog-
nized to provide additional health benefits that may reduce
disease risks and/or promote optimal health. Functional foods
include: (1) conventional foods or whole foods, (2) modified
foods, (3) medical foods, and (4) foods for special dietary use.
According to Functional Food Science in Europe, Functional
foods are those satisfactorily demonstrated to affect beneficially
one or more target functions in the body, beyond adequate
nutritional effects, in a way that is relevant to either an improved
state of health and well-being and/or reduction of risk of disease
when consumed as part of normal food pattern. Thus, func-
tional foods include: a natural, unmodified food; a food in
which one of the components has been enhanced through spe-
cial growing conditions, breeding, or biotechnological means; a
food to which a component has been added to provide benefits;
a food from which a component has been removed by techno-
logical or biotechnological means so that the food provides
benefits not otherwise available; a food in which a component
has been replaced by an alternative component with favorable
properties; a food in which a component has been modified by
enzymatic, chemical, or technological means to provide a bene-
fit; a food in which the bioavailability of a component has been
modified and combination of any of the above.
In Japan, FOSHU refers to foods consumed as part of an
ordinary diet, containing functional ingredients and exerting
health or physiological effect. FOSHU requires that safety
assessment, health functions, and claims must be approved
by the Ministry of Health, Labour, and Welfare. Functional
foods are also required to have three other fundamental fea-
tures: (1) nutritional functions, (2) sensory functions, and (3)
physiological functions such as regulation of biorhythms, the
nervous system, the immune system, and body defense beyond
nutrient functions. As of April 2011, in Japan, 955 products
were approved as FOSHU.
Based on extensive literature review and expert consultation,
A functional food is, or appears similar to be, a conventional
food. It is part of a standard diet and is consumed on a regular
basis, in normal quantities. It has proven health benefits that
reduce the risk of specific chronic diseases or beneficially affect
target functions beyond its basic nutritional functions.
Even though there are slight variations, the following are
reflected on definitions given by different sources: (1) func-
tional foods should be consumed as part of regular safe diet
not as drugs; (2) they are required to bear functionally bioac-
tive adequate ingredient(s) beyond basic nutrients; (3) diet
with its functionally bioactive ingredient(s) should impact
positive health in an individual by either (i) decreasing occur-
rences of disease(s), (ii) decreasing disease-causing factor(s), or
(iii) promoting positive human physiological functions for
optimal health such as by maintaining body homeostasis
and/or through bolstering body immune systems; (4) they
should improve quality of life; and (5) if there is no history
of consumption as a regular diet, they must pass a strict regu-
latory evaluation process for their safety and for beneficial
claim(s) declared substantiated with sound scientific evidence.
Other Concepts Related to Functional Foods
Nutraceuticals
These are distinct from functional foods and defined as a
product isolated or purified from foods generally sold in
medicinal forms (i.e., as pills, powder, syrups, or other medic-
inal forms) that have specific health benefits.
Dietary Supplements
These are products other than tobacco intended to supplement
diet, which contain one or more of the following dietary ingre-
dients: vitamin, mineral, herb or other botanical, amino acid,
concentrate, metabolite, constituent, extract, or combinations
of these ingredients; they are ingested in pill, capsule, tablet, or
liquid form; they are not represented for use as conventional
food or as a sole item of diet.
 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview
Novel Foods
These are defined as raw foods or food materials with no
history of safe use, for example, genetically modified foods;
foods produced by algae, fungi, or microorganisms; foods
isolated from plants and animals without a history of safe
use; and foods that have been processed where the process
has dramatically changed the food.
Bioavailability
This refers to a fraction of a given compound and/or its metab-
olite that reaches systemic circulation and involves gastrointes-
tinal digestion, absorption, metabolism, and tissue
distribution. Bioavailability encompasses bioaccessibility and
bioactivity.
Bioaccessibility
This refers to the quantity or fraction of a bioactive that is
released from a food matrix in the gastrointestinal tract and
becomes available for absorption. It includes digestive trans-
formations, absorption/assimilation into intestinal epithelium
cells, and also the presystemic metabolism (both intestinal and
hepatic).
Bioactivity
This refers to a specific response, biomarker, or effect observed
upon exposure to the bioactive substance. It includes tissue
uptake and the subsequent physiological response.
Recognized Bioactive Compounds in Functional Food
Phenolics (Polyphenolics)
Phenolics are ubiquitous in plants and more than 8000 com-
pounds are known. Dietary phenolics can be broadly divided
into flavonoids (anthocyanins, flavonols, flavones, flavanones,
isoflavones, flavan-3-ols, condensed tannins or proanthocya-
nidins, lignans, chalcones, coumarins, coumestans, and aur-
one) and nonflavonoids (simple phenols, phenolic acids
(hydroxybenzoates and hydroxycinnamates), and stilbenes)
(Figure 1). Flavonoids are the largest group and more than
6000 compounds have been identified. Hydrolyzable tannins
are derivatives of gallic acids. They exert deleterious effects and
may be found at trace levels in some edible foods. Flavonoids
often exist as glycosides with sugar molecules attached often at
position 3, sometimes at 5, 7, 40 , 30 , and 50 and hydroxyls at 40 ,
5, and 7 on the flavonoid basic structure. Flavonoids without
sugars are called aglycones.
Phenolic acids and various flavonoids in whole grains and
bran fractions (pericarp, aleurone layer, testa, and germ) are
extensively documented. Bran is removed during cereal grain
milling for production of refined flours. Among phenolic
acids, ferulic acids are dominant (about 80% to 90%) in cereal
grains. Sorghum is known to have unique 3-
deoxyanthocyanins. Other unique phenolics documented in
grains are alkylresorcinols in rye, wheat, triticale, and barley,
avenanthramides in oats, and g-oryzanols (ferulic cid esters of
3
phytosterols and triterpene alcohols) in rice (Figure 2). Alkyl-
resorcinol metabolites are currently studied for use as
biomarkers of whole-grain intake. Bran also contains vitamins,
phytosterols, minerals, proteins, lipids, carotenoids, tocols,
and phytic acids. Consumption of grain foods along with
their bran is beneficial due to the presence of various bioactive
compounds. Functional roles documented for phenolics
include: bolstering of cellular antioxidant defenses, protection
from cardiovascular disease (CVD) risks, brain wellness, anti-
neurodegenerative, antiulcer, anticarcinogenic, antimutagenic,
anti-inflammatory, antimicrobial properties; and improve-
ment of eye health and vision, muscle performance, and
immune responses.
Phenolics are generally partially bioavailable and their
properties are influenced by their structures and solubility.
Most soluble phenolics are found unbound, whereas insolu-
bles are found bound to cell wall polysaccharides or proteins
where they form stable complexes. Insoluble phenolics are not
absorbed in the small intestine, but they are fermented by
colon microbiota yielding largely ferulic acid followed by
p-coumaric acid. Within cells, phenolics suppress reactive-
oxygen (ROS) and nitrogen species (RNS) by donating hydro-
gen atoms or electron, quenching free radicals, and activating
endogenous antioxidant enzymes. They also chelate metal pro-
oxidants. Oxidative stress can happen when generation of
oxidants exceeds the natural antioxidant systems. Oxidative
stress causes damages to DNA, RNA, lipids, proteins, and car-
bohydrates and leads to different types of chronic diseases. In
cereal grains, anti-oxidant potentials follow pigmented grains
(tannin sorghums, black rice, brown and black sorghums, teff,
millets, blue maize) > non-pigmented grains (white-sorghum,
wheat, rice). Even though, phenolics are shown to suppress
cancers in in vitro in cells and in vivo in animal models, studies
in human intervention are less clear. The synergetic effects are
believed to have a role. Phenolics are also used as antimicrobial
food preservatives, food colorants, and flavor imparters. Tan-
nins, despite having high antioxidant activity, are antinutri-
tional and inhibit digestive enzymes and mineral absorptions.
Phytoestrogens
Phytoestrogens are nonsteroidal plant compounds with struc-
tures and biological activities similar to steroid hormone estro-
gen, 17b-estradiol (Figure 3). Functional roles documented
include: influence on growth and functioning of female and
male reproductive tissues; maintenance of skeletal and central
nervous systems; cardio-protection; suppression of colon,
breast, and prostate cancers and skin aging; and relief from
menopausal symptoms. At low concentration, they act like
estrogens (agonists) and at high doses as blockers (antago-
nists). Phytoestrogens are potential endocrine disrupters and
become cytotoxic at high dose. Groups of compounds recog-
nized as phytoestrogens are lignans, isoflavones, coumestans,
and stilbenes.
Lignans
These are 18-carbon skeleton compounds composed of two
phenylpropanoid units with various functional groups con-
nected to benzene rings (Figure 4). Secoisolariciresinol,
4 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview

OH O OH
Derivatives of simple phenol C Derivatives of hydroxybenzoic acid
R1 Phenol: R1=R2=R3=H Gallic acid: R1=H, R2=R3=R4=OH
R1 Gentisic acid: R1=R4=OH, R2=R3=H
Catechol: R1=OH, R2=R3=H
Salicylic acid: R1=OH, R2=R3=R4=H
R2 R3 Resorcinol: R1=R3=H, R2=OH R4 p-(OH) Benzoic acid: R =R =R =H, R =OH
R2 1 2 4 3
Phloroglucinol: R1=H, R2=R3=OH
Protocatechuic acid: R1=R4=H, R2=R3=OH
OH R3
Syringic acid: R1=H, R2=R4=OCH3, R3=OH
C Derivatives of hydroxycinnamic acid Vanillic acid: R1=R4=H, R3=OH, R2=OCH3
R1 O Cinnamic acid: R1=R2=R3=R4=H
o- Coumaric acid: R1=OH, R2=R3=R4=H
m- Coumaric acid: R1=R3=R4=H, R2=OH
R2 R4
p- Coumaric acid: R1=R2=R4=H, R3=OH
R3 Caffeic acid: R1=R2=H, R3=R4=OH
Ferulic acid: R1=R2=H, R3= OH, R4 =OCH3
Sinapic acid: R1=H, R2=OCH3, R3=OH, R4=OCH3
Phenolic acids

Stilbene

3`
2`
4`
8
1 O
7 O 2 5` O O
6` Flavan-4-o1
6 3 OH OH
5 4 OH
Flavonoid basic structure Flavan-3-o1 Flavonol
O O
OH
O
OH
+ Flavone Flavanone
O O
HO O
O O
OH
Anthocyanidin Isoflavone O
OH O Aurone
O O

O O O Lignan
Chalcone Coumarin
Coumestan
O

Basic structures of main class flavonoids and related structures (aurone, lignan)
OH
OH

HO O
R OH n:0 to many
OH OH
OH
OH
HO O
HO +O
OH
OH
OH
OH
OH HO O
Sorghum 3-deoxyanthocyanidins
(Apigenindin, R=H and Luteolinidin, R=OH) OH
OH Proanthocyanidins

Figure 1 Structures of phenolics.

 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview 5

R1 O
OH R2
N
H
O C OH R3
HO R Avenanthramides (more than 35 forms are identified)
Alkylresorcinols (R = alkyl chain, Avenanthramide 2p, R1=R3=OH, R2=H
odd carbon C15 to C27) Avenanthramide 2f, R1=R3=OH, R2=OCH3
Avenanthramide 2c, R1=R2=R3=OH
O
CH3O C OR
O
CH3O C O
HO
Gamma-oryzanol (ferulic acid esters,
HO
R = phytosterols or triterpene alcohols) Gamma-oryzanol (campesteryl ferulate)

Figure 2 Structures of alkylresorcinols, avenanthramides, and gamma-oryzanols.

OH OH O
OH
H H

H H H H H H
HO HO HO
17 beta-Estradiol Estriol Estrone

Figure 3 Structures of mammalian endogenous estrogens.

HO OH
O OH
H
H CH2O
CH3O CH2OH CH3O HO OH
H OH
H
HO CH2O O
HO CH2OH
OH
OH
OCH3 OCH3
Secoisolariciresinol diglycoside
Secoisolariciresinol OH
OH
O
H O
O H
CH3O O
O HO
O O
HO Sesamolin
O O
Matairesinol O OH O
O
O H
OCH3 H O
O O
OH Sesamol O
Sesamin O
H H
HO OH HO
O
OH
Enterolactone O
Enterodiol

OH OH

Figure 4 Structures of common lignans.

6 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview
matairesinol, sesamin, and sesamolin are typical examples.
When two glucose molecules are attached to the hydroxyl
groups of the propanol chain of secoisolariciresinol, the com-
pound is called secoisolariciresinol diglucoside (SDG). Lig-
nans have higher antioxidant potential than vitamin E.
Lignans are implicated to have potential use in hormone
replacement therapy, to maintain good cognitive function in
postmenopausal women, and to have anticancer activities
(breast, prostate, colon, and skin cancers). They may contrib-
ute to the reduction of hypercholesterolemia, atherosclerosis,
and diabetes. SDG is converted by bacteria in the human
colons to mammalian lignans enterodiol and enterolactone,
which are also antioxidants. Enterodiol and enterolactone are
purported to reduce growth of cancer tumors, especially
hormone-sensitive ones. Grain sources of lignans include oil-
seeds (flax, rapeseed, and sesame), whole-grain cereals (wheat,
oats, rye, barley, and millets), and legumes (soybean). Flaxseed
is the richest source of SDG and also contains other lignans
(matairesinol, pinoresinol, lariciresinol, and isolariciresinol).
Baked products and salad dressings fortified with flax and
sesame seeds can be considered as functional foods rich in
lignans
Isoflavones
Isoflavones are often found as glycosides. Common isofla-
vones include daidzin, genistin, biochanin A, and formonone-
tin (Figure 5). The glycosides are converted to aglycones by gut
microflora. Among isoflavones, studies with genistein on
breast cancer cell lines show cell proliferation at low doses
and inhibition at high concentrations. Isoflavones are also
recognized as mild antioxidants.
Isoflavones are exclusively documented in leguminosae of
which soybean is the richest source. Other sources are kidney,
navy, pinto, red, small white, and mung beans and chickpeas,
split peas, peanuts, sunflower seed, and walnut. Raw soybeans
can contain 1.24.2 mg g1 dry weight isoflavones, while high-
protein soy flour may contain up to 1.11.4 mg g1 dry weight.
Diets rich in soybeans are implicated to improve cognitive abil-
ities, prevent osteoporosis and various cancers, improve immune
R1 O R2 Isoflavones
R1 R2
Geinstein OH OH
HO O Daidzin H OH
Biochanin A OH OCH3
Formononetin H OCH3
Figure 5 Structures of isoflavones.
OH
O
HO O HO
O
Coumestrol
Figure 6 Structures of coumestrol and 40 -methoxycoumestrol.
function and maintenance of menopausal health in women, and
also suppress CVD by lowering total cholesterol and LDL
cholesterol and raising HDL in the blood vessels. However, the
isoflavone content of soybean products varies with processing.
Water- and alcohol-extracted soybean protein concentrate and
soybean oils have insignificant levels of isoflavones.
Coumestans
Coumestans are less abundant than isoflavones in human
diets. They are known to have stronger estrogenic activity
than isoflavones. Typical examples are coumestrol and 40 -
methoxycoumestrol (Figure 6). Coumestrol and genistein have
higher binding affinities to ERb than other phytoestrogens.
Coumestrol has been reported to inhibit bone resorption and
stimulate bone mineralization by in vitro studies. Among grains,
chief sources are soy sprouts, mung bean, and sunflower seeds.
Stilbenes
Stilbenes are C6C2C6 compounds with H,OH, and OCH3
as substituents on their two benzene (C6) rings (Figure 7).
Stilbenes are known to be anticarcinogenic, antioxidative,
antimutagenic, antitumor, and antiosteoporotic. Diets rich in
resveratrol, which exists in both trans and cis isomers of the
stilbene, are purported to prevent CVD, neurodegenerative dis-
orders such as Parkinsons and Alzheimers disease, and various
cancers. Resveratrol has stronger antioxidant activity than propyl
gallate, vanillin, phenol, 2,6-di-tert-butyl-p-hydroxytoluene
(BHT), and a-tocopherol. Among grains, stilbenes are found in
Fabaceae and Poaceae. Other good sources are peanuts, grape
skins, and notably some sorghum varieties.
Phytosterols
Phytosterols are natural components of human diets, are cho-
lesterol-like and lipophilic, and comprise sterols and stanols.
Stanols are hydrogenated products of sterols. Of more than
250 sterols and stanols known to exist in nature, only six
(campesterol, campestanol, stigmasterol, sitosterol, sitostanol,
and brassicasterol) are dominant in seed oils (rapeseed,
soybean, corn, and sunflower oils), other grains (corn, rye,
R HO
R OH
R HO
Stilbenes,R = H, OH or OCH3 Resveratrol
Figure 7 Structures of stilbenes.
O OCH3
O O
4`-M ethoxycoumestrol
 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview 7

R 26
18 R 20
12 20 23
11 21 20 22 25 21 24
17 16 21 20 22 23 24 25 26 23 26 25
19 13 24 22 27
1 9
H 14 15
2 27 Brassicasterol
27
10 8
Cholesterol/stanol Sitosterol/stanol
5 7
HO 3
4 6 18 R20
12
Sterols 11 21 20 22 23 25 26
17 16 21 20 22 23 25 26
19 24
1 9
H 13
14 15
24
2 27
27
10 8 Stigmasterol/stanol
5
Campesterol/stanol
7
HO 3
4 6
(a) Stanols

C O
Sitosteryl stearate
O
H
H3CO

HO CH CHCOO
Sitostanyl ferulate
H
CH2OH
O
OH O
HO Steryl glycoside (SG)
OH
O
C O CH H
2
O
OH O
HO Acylated steryl glycoside (ASG)
(b) OH
Figure 8 (a) Structure of plant stanols and plant sterols. (b) Conjugated structure of plant stanols and plant sterols.

wheat, barley, millets, rice, oats, and peanuts), and tree oils
(Figure 8(a)). In plant tissues, sterols and stanols can exist as
conjugates often esterified with fatty acids, oleic, or linoleic in
five forms (Figure 8(b)).
Plant sterols and stanols are known to attenuate absorption
of low-density lipoprotein cholesterol (LDLC), a risk factor for
premature atherosclerosis and CVD. Consumption of 2 g d1
of plant sterols or stanols as part of diet is known to lower
LDLC plasma levels by about 10%. Consumption of low cho-
lesterol and saturated fat along with high stanols or sterols can
reduce LDL by 20%. Statin medication along with sterols or
stanols is more effective in lowering LDL than doubling statin
dose. Ranking toward blocking of cholesterol absorption is as
follows: free-form stanols > stanol esters > free-form sterols >
sterol esters due to structural and metabolic reaction differ-
ences toward gut microorganisms. The precise mechanisms of
cholesterol-lowering effect by individual compounds in
humans remain largely unknown. Plant sterols/stanols are
also implicated to have mild anti-inflammatory effects and
are not toxic at physiological concentrations.
Carotenoids
Carotenoids are lipid-soluble tetraterpene hydrophobic
compounds, which impart yellow, orange, or red colors, and
found associated with lipid portions in human cells. Carotenoids
are divided into two groups (Figure 9): carotenes (hydrocarbons)
and xanthophylls (oxygenated carotenes). Carotenoids bear con-
jugated double bonds in their structures, which are responsible
for color formation and reactive oxidant stabilization. Caroten-
oids are antioxidants, and functional foods processed from carot-
enoids are helpful for prevention of CVD and cancers. b-carotene,
a-carotene, and b-cryptoxanthin are provitamin A compounds.
8 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview

CH3 CH3 H3C

H3C CH3 Beta-carotene
Examples of
carotenes
H3C CH3
CH3 CH3
CH3
CH3 CH3 H3C
Alpha-carotene
H3 C CH3
H

H3C CH3
CH3 CH3
CH3
CH3 CH3 H3C
H3C CH3 Beta-cryptoxanthin

H3C CH3 Examples of

HO CH3 CH3
CH3 xanthophylls
CH3 CH3 H3C OH
CH3 Zeaxanthin*
H3C

H3 C CH3
HO CH3 CH3
CH3
CH3 CH3 H3C OH
CH3 Lutein*
H3C

H3C CH3
HO CH3 CH3
CH3

CH3 CH3 O Retinyl palmitate

H3C CH3
Vitamin A
O compounds

CH3 H3C

CH3 CH3 O
H3C CH3
OH Retinoic acid

CH3
CH3 CH3 O CH3 CH3
H3C CH3 H3C CH3
H OH
Retinal Retinol
CH3 CH3

Figure 9 Structures of carotenoids (*not provitamin A).

Of more than 700 carotenoids identified in nature, 24 occur
commonly in human foods and 40 have been detected in
human milk, serum, and tissues.
Among grains, chief sources are yellow maize, durum
wheat, einkorn, and golden rice. Carotenoid contents can
vary depending on varieties, cultivation conditions, geographic
locations, part of plant utilized, maturity stage, food proces-
sing, and storage conditions used. Heating, cooking, and
mechanical disruption and the presence of fats/oils can
enhance bioavailability of carotenoids. Carotenoid activities
can be destroyed by isomerization and oxidative reactions.
Lutein and zeaxanthin are not synthesized by humans. They
can be supplied from grains (yellow maize, durum wheat,
golden rice, and einkorn), vegetables (basil, parsley, spinach,
kale, leek, and red pepper), yellow egg yolks, and human milk.
Along with their metabolite meso-zeaxanthin, both are
protective against age-related macular degeneration and cata-
racts, a leading cause of eye blindness. Cereal-based lutein and
zeaxanthin-containing functional foods processed from yellow
maize, durum wheat, golden rice, einkorn, and maize bran
include high-lutein bread, high-lutein cookies, high-lutein
muffins, corn tortilla, corn chips, and extruded products.
Omega (v)-3 Fatty Acids
Omega-3 fatty acids are long-chain polyunsaturated fatty acids
(PUFA) (C18-C22) commonly designated as o-3 fatty acids
(Figure 10). The o-3 and o-6 fatty acids are essential nutrients.
Alpha-linolenic acids (ALA, o-3) derived from plant sources
are precursors for physiologically important longer-chain
 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview
ALA (C18:3 9,12,15)
EPA (C20:5 5,8,11,14,17)
DHA (C22:6 4,7,10,13,16,19)
Figure 10 Omega-3 fatty acids: ALA, EPA, and DHA.
PUFAs: eicosapentaenoic (EPA, o-3) and docosahexaenoic acid
(DHA, o-3).
Omega-3 fatty acids are components of cell membranes.
Their documented health benefits include suppressions of
CVD, cancer, type 2 diabetes, depression and stress, inflamma-
tory diseases, and (6) Alzheimers disease. They are also bene-
ficial in infant nutrition for normal brain, eye, and mammalian
nervous system development and function. Omega-3 fatty
acids are available as supplements from pharmaceuticals.
However, consumers prefer o-3 fatty acids from diets. In this
respect, there is high opportunity for o-3 fatty acid-containing
functional foods. The main source of ALA is flaxseed. Other
sources are canola, soybean oils, and walnuts. In flaxseed,
bioavailability of ALA is as follows: oil > milled > whole
flaxseed. Despite fish oil being a major source of EPA and
DHA, it is sensitive to autoxidation, and there is also a possi-
bility for methyl mercury toxicity since fish has potential for
contamination with this toxin. Where consumption of fish oil
is limited and methyl mercury toxicity is a possibility, EPA and
DHA are required to be supplied from ALA even though con-
version of ALA to EPA and DHA is known to be inefficient.
Baked foods like bread and injera and various cereal grain
snacks processed by incorporating flaxseed oils and/or milled
flaxseed flours are an alternative way of supplying o-3
fatty acids.
Gluten-Free Foods
In some genetically predisposed individuals, gluten proteins
from wheat, rye, barley, and triticale when ingested will lead to
celiac disease (CD). Oats are also not considered free, because
of potential contamination and the possibility that some peo-
ple who are intolerant to gluten are not immune to it. CD
involves interactions between genetic susceptibility (HLA-
DQ2 and HLA-DQ8 alleles), gluten proteins, and an
immunologic response that causes small intestinal mucosal
damage characterized by complete loss of absorptive villi to
maldigestion and malabsorption of nutrients. Such effects lead
to chronic diseases like skin lesions, anemia, diarrhea, weight
loss, osteoporosis, fatigue, infertility, and cancer. About 1% of
the world population is estimated to be gluten-intolerant.
There is no drug for CD, and the remedy is a complete adher-
ence to a gluten-free diet (GFD). The Codex Alimentarius Com-
mission and FDA have set standards for GFD to be
<20 mg kg1 gluten. Gluten proteins are vital for production
of aerated baked products to which consumers are attracted.
The use of baked products from gluten-free cereal grains (teff,
maize, rice, sorghum, and the millets) particularly those pro-
cessed from whole grains is a remedy for CD. Teff grain is one
9
O
C OH
O
C OH
O
C OH
such best candidate because it is always consumed as whole
grain often fermented as injera with insignificant antinutrients
in it and in many respects it supplies better nutrients for celiac
patients. Other approaches to gluten-free bread quality
improvements are the use of gluten-free flours and starches,
incorporation of nutritional ingredients, and use of structure
modifier additives and processing technologies that will
change the allergenic glutens.
Other bioactive compounds to be noted include tocols
(tocopherols and tocotrienols) from cereal germ and aleurone
layers; betaine and choline from wheat, barley and rye germ,
and brans; and benzoxazinoids from whole-grain wheat and
rye. Some bioactive compounds known for their antinutri-
tional effects tend to portray beneficial effects under specific
conditions. They include glucosinolates from rapeseed, phytic
acids from germ and aleurone layers of grains, and lectins
(hemagglutinins) from legumes and whole cereal grains.
Conclusions
Opportunities for grain-based functional foods are high
because grains are sources of many bioactive compounds that
are located in specific anatomical structures. However, bio-
availability can be influenced by many factors including
forms of existence, processing technology, and food matrices
that prevail during consumption. Functional foods are recog-
nized to be consumed as varied regular diets. However, the
absence of a definition that has received global consensus
potentially poses various risks such as inaccurate health claims,
adverse reactions, poor quality control during manufacturing,
allergic reactions, and carcinogenicity. Novel foods in particu-
lar may pose unprecedented risks because they do not have a
long history as diets to populations of different genetic back-
ground. A distinction between novel and functional foods may
help to reduce such effects. Misuse of functional foods can
lead to adverse effects such as anti-nutritional, thyroid toxicity,
drug interactions, allergic reactions and geno-toxicity/
carcinogenicity.
A given diet is believed to bear more than 25 000 bioactive
constituents. Minimum dietary intake that would lead to desir-
able health effect and maximum intake to avoid adverse effects
are yet not available for most bioactive compounds. A distinc-
tion is yet to be clearly established for most bioactive com-
pounds whether a desirable outcome is caused by a single
compound or by synergic effects. Since they are consumed as
regular varied diets for health promotion, designing functional
foods for synergic effects of bioactive compounds seems ben-
eficial. While designing for an optimum supply for those
health effects is established, caution is also required not to
10 FOOD GRAINS AND WELL-BEING | Functional Foods: Overview
affect other components used to be served in a given regular
diet. With an accurate definition in place, functional foods can
be better regulated to improve health and well-being.
Exercises for Revision
Provide three features that you think useful to describe
functional foods.
Describe how functional foods are different from dietary
supplements, prescribed drugs, traditional medicines, and
nutraceuticals.
List various grain-based bioactive compounds.
Provide factors that may influence bioaccessibility, bio-
availability, and bioactivity of bioactive compounds.
List potential safe diets for celiac patients.
Name indigenous foods consumed in your communities
believed to have health-promoting effects.
Exercises for Readers to Explore the Topic Further
Explore the possibility of extending the earlier lists of bio-
active compounds in various grains.
What metabolites are produced from the partial degrada-
tion of some of the bioactives listed? Find out if these are
still active.
Investigate what innovative processing technologies might
provide optimum utilization of grain bioactive com-
pounds. Could these technologies also ensure that food
safety issues are addressed because bioactive compounds
are located in unique grain anatomical parts?
Research the possibilities in plant breeding to enhance the
presence and availability of bioactives.
Investigate the possible exploitation of indigenous foods
relevant to your region. Their consumption as safe diets has
been known for ages, so their potential to grow in the
functional foods spectrum should be considerable.
See also: Appendix 3: Grains, Foods, and Ingredients Suiting
Gluten-Free Diets for Celiac Disease; Bioactives and Toxins:
Bioactives: Antioxidants; Bioactive Compounds in Wheat Bran; The
Antinutritional Components of Grains; Carbohydrates: Glycemic
Index, Glycemic Load, and Their Health Benefits; Grains other than
Cereals: Non-starch Polysaccharides; Food Grains and the
Consumer: Cultural Differences in Processing and Consumption;
Fortification of Grain-Based Foods; Genetically Modified Grains and the
Consumer; Grains and Health: Misinformation and Misconceptions;
Grains and Health; Labelling of Grain-Based Foods; Food Grains and
Well-being: Functional Foods: Dietary Fibers, Prebiotics, Probiotics,
and Synbiotics; Food Grains: Intolerance, Allergy and
Diseases: Celiac Disease; Cereal Allergens; The Gluten-Free Diet;
The Basics: Grain: Morphology of Internal Structure.
Further Reading
Bagchi D (ed.) (2014) Nutraceutical and Functional Food Regulations in the United
States and Around the World. NW, USA: Elsevier Inc.
Capriles VD and Areas JAG (2014) Novel approaches in gluten-free bread making:
interface between food science, nutrition, and health. Comp. Rev. Food Sci. Food
Saf. 13: 871890.
Doyon M and Labrecque J (2008) Functional foods: a conceptual definition. Br. Food J.
110(11): 11331149.
Dykes L and Rooney LW (2007) Phenolic compounds in cereal grains and their health
benefits. Cereal Foods World 52: 105111.
Fraga CG (ed.) (2010) Plant Phenolics and Human Health: Biochemistry, Nutrition, and
Pharmacology. Hoboken, NJ, USA: John Wiley & Sons, Inc.
Gogus U and Smith C (2010) n-3 Omega fatty acids: a review of current knowledge. Int.
J. Food Sci. Technol. 45: 417436.
Gylling H, Plat J, Turley S, Ginsberg HN, Ellegard L, Jessup W, Jones PJ, Lutjohann D,
Maerz W, Masana L, Silbernagel G, Staels B, Boren J, Catapano AL, De Backer G,
Deanfield J, Descamps OS, Kovanen PT, Riccardi G, Tokgozoglu L, and
Chapman MJ (2014) Plant sterols and plant stanols in the management of
dyslipidaemia and prevention of cardiovascular disease. Atherosclerosis
232: 346360.
Hasler CM and Brown AC (2009) Position of the American dietetic association:
functional foods. J. Am. Diet. Assoc. 109: 735746.
Howlett J (2008) Functional Foods: From Science to Health and Claims. Belgium: ILSI
Europe.
Malla S, Hobbs J, and Sogah EK, (2013) Functional Foods and Natural Health Products
Regulations in Canada and Around the World: Nutrition Labels and Health Claims.
Report Prepared for the Canadian Agricultural Innovation and Regulation Network
(CAIRN), Canada
Mohamed S (2014) Functional foods against metabolic syndrome (obesity, diabetes,
hypertension and dyslipidemia) and cardiovascular disease. Trend Food Sci.
Technol. 35: 114128.
Ndolo VU and Beta T (2014) Comparative studies on composition and distribution of
phenolic acids in cereal grain botanical fractions. Cereal Chem. 91(5): 522530.
Peterson J, Dwyer J, Adlercreutz H, Scalbert A, Jacques P, and McCullough MJ (2010)
Dietary lignans: physiology and potential for cardiovascular disease risk reduction.
Nutr. Rev. 68(10): 571603.
Sajilata MG, Singhal RS, and Kamat MY (2008) The carotenoid pigment zeaxanthina
review. Comp. Rev. Food Sci. Food Saf. 7: 2949.
Taylor JRN, Belton PS, Beta T, and Duodu KG (2014) Review: Increasing the utilization
of sorghum, millets and pseudocereals: Developments in the science of their
phenolic phytochemicals, biofortification and protein functionality. J. Cereal Sci.
59: 257275.
Relevant Websites
http://www.biomedcentral.com/content/supplementary/1475-2891-9-3-S1.PDF
University of Oslo.
http://www.glnc.org.au/wp-content/themes/glnctheme/images/home.png
http://www.google.co.bw/url?
http://www.nutritionj.com/content/9/1/3.
 Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics
G Bultosa, Botswana College of Agriculture, Gaborone, Botswana; Haramaya University, Dire Dawa, Ethiopia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Dietary fiber (DF) and whole grain functional foods.
Prebiotic, probiotic, and synbiotic functional foods.
Metabolic syndrome-related diseases and cancers.
Lactose intolerance.
Learning Objectives
To achieve understanding of the definitions of DFs, whole
grain foods, prebiotics, probiotics, and synbiotics.
To achieve understanding of DF and prebiotic grain sources.
To impart principles of DFs, prebiotics, probiotics, and
synbiotics with respect to the processing of functional foods.
To explain linkages of the intake of DFs, prebiotics, probio-
tics, and synbiotics to health.
Introduction
Functional foods processed from DFs are linked to various
health benefits. Soluble fibers and b-glucans in particular are
known to modulate hyperlipidemia, hyperinsulinemia,
hyperglycemia, and obesity, which are risk factors for chronic
diseases including cardiovascular diseases (CVDs), diabetes
mellitus, and cancers. Soluble fibers are rapidly fermented by
gut microbiota. Adequate intake (AI) of insoluble fibers is also
linked to the reduction of colon cancers and improved gastro-
intestinal health (GIH), reduced CVD, and obesity. Cereal
grain fibers and legume cotyledons are included in bread,
extruded products, cakes, cookies, and fermented and other
ready-to-eat breakfast cereal (RTE) products. Consumption of
whole grain foods is encouraged for AI of DF. Health-
promoting effects of DF are imparted by nonstarch carbohy-
drates, resistant starches, and closely associated/attached
compounds. Those DFs that selectively promote growth and/
or activities of probiotics and promote host health are pre-
biotics. Arabinoxylans, fructans, inulin, resistant starches, b-
glucans, and a-galactooligosaccharides are fermented by pro-
biotics to short-chain fatty acids (SCFAs): butyrate, propionate,
and acetate that are absorbed in the colon to furnish additional
energy for the host. Prebiotics are also known to improve the
absorption of calcium and magnesium and to suppress colon
cancers and absorption of triglycerides and cholesterol. Cellu-
lose and water-insoluble arabinoxylans are major contributors
to laxative effects and fecal bulking. Barley, wheat, rye, oats,
triticale, maize, einkorn, chickpea, and lentils are notable
sources of prebiotics. Intake of prebiotics is also achieved
through consumption of whole grains like teff, millets,
sorghums, and brown rice. Dietary intake of probiotic bacteria
mainly from the genera of Lactobacillus and Bifidobacterium is
linked to improved immune capacity and GI health and to
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00245-X
suppressing various diseases including colon cancers, allergy,
asthma, celiac disease, HIV infection, inflammatory bowel
diseases, hypertension, gastrointestinal pathogen infections,
peptic ulcer, diarrhea, obesity, necrotizing enterocolitis, rheu-
matoid arthritis, and liver diseases. In this article, the nature
and sources of DFs, prebiotics, probiotics, and synbiotics for
functional foods are described with emphasis on grains.
Dietary Fibers and Whole Grains
Dietary Fibers
DFs are a structurally diverse group of compounds. Various
regulatory bodies and professional societies American Asso-
ciation of Cereal Chemists International (AACCI), Institute of
Medicine (IOM), Food Standards Australia and New Zealand
(FSANZ), FAO/WHO Codex Alimentarius Commission, Euro-
pean Food Safety Authority, and Health Canada have offered
definitions for DF primarily based on the physiological roles
and analytical methods used to isolate and quantify them.
According to Codex, DFs are carbohydrate polymers with 10
or more monomeric units (decision on whether to include
carbohydrates of 39 monomeric units should be left up to
national authorities), which are not hydrolyzed by the endog-
enous enzymes in the small intestine of humans and comprise
the following:
(1) Edible carbohydrate polymers naturally occurring in the
food as consumed. When derived from a plant origin, DF
may include fractions of lignin and/or other compounds
protein fractions, phenolic compounds, waxes, saponins,
phytates, cutin, vitamins, phytosterols, minerals, etc.
when such compounds are associated with the polysac-
charides in the plant cell walls and extracted together.
(2) Carbohydrate polymers, which have been obtained from
food raw materials by physical, enzymatic, or chemical
means and which have been shown to have a beneficial
physiological effect to health as demonstrated by generally
accepted scientific evidence to competent authorities.
(3) Synthetic carbohydrate polymers, which have been shown
to have a beneficial physiological effect to health as dem-
onstrated by generally accepted scientific evidence to com-
petent authorities.
The Codex definition is expected to be used in all jurisdictions.
Currently, there is wider consensus that the AACCI definition is
reconciling with Codex definition. AACCI defines DF as edible
parts of plants or analogous carbohydrates that are resistant to
digestion and absorption in the human small intestine with
complete or partial fermentation in the large intestine. DFs
include polysaccharides, oligosaccharides, lignin, and associated
plant substances. DFs promote beneficial physiological effects
including laxation and/or blood cholesterol and glucose attenu-
ations. DFs thus comprise the following:
11
12 FOOD GRAINS AND WELL-BEING | Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics
i. Nonstarch polysaccharides and resistant oligosaccha-
rides: cellulose, b-glucans, hemicellulose (arabinoxy-
lans and arabinogalactans), polyfructoses (inulin and
oligofructans), galactooligosaccharides, gums,
mucilages, and pectins
ii. Analogous carbohydrates (indigestible dextrins/resis-
tant maltodextrins from corn and other sources and
resistant potato dextrins), synthesized carbohydrate
compounds (polydextrose, methylcellulose, and
hydroxypropyl methylcellulose), and resistant starches
iii. Lignins
iv. Substances associated with the nonstarch polysaccha-
rides and lignin complex in plants (waxes, phytates,
cutins, saponins, suberin, polyphenols, tannins,
vitamins, minerals, alkylresorcinols, carotenoids, beta-
ine, etc.)
The classification of DFs as soluble (gums, oligosaccharides,
pectins, water-soluble arabinoxylans, b-glucans, galactoman-
nans, psyllium fibers, and alginates) and insoluble (cellulose,
lignins, and water-insoluble arabinoxylans) also continues to be
in use. DFs for human nutrition are largely derived from cereal
grains 44%, vegetables 21%, fruits 13%, and legumes 10%. The
outer pericarp of cereal grains, endosperm cell walls, seed coat
and cotyledons of legumes, psyllium husks, and fruit and vege-
table skins are rich sources of DF. Diets rich in bran (pericarp,
aleurone layer, and testa) and germ are known to have high
antioxidant activities than diets based on refined white flours
largely due to associated DFphenolics, tocols, and carotenoid
antioxidants. Functional foods containing DFs are currently
widely available in the form of breads; extruded products like
pasta, cakes, and cookies; and fermented and other RTE prod-
ucts. Although incorporation of DF to some levels has produced
products comparable to those processed from refined flours,
increased DF contents have been associated with reduced con-
sumer acceptance (color, texture, and taste). Currently, improve-
ments being researched include the use of enzymes and other
physicochemical means to modify fiber structures and incorpo-
ration of fibers from fruits and vegetable skins.
The beneficial physiological effects conferred by AI of DF
include attenuation of blood cholesterol and blood sugars,
suppression of CVD diseases (coronary heart disease, stroke,
and hypertension), weight control, improved insulin sensitiv-
ity and suppression of type 2 diabetes mellitus, improvement
of GIH, improved laxation, suppression of cancers such as
colon and breast cancers, improved immunity, and contribu-
tion to the overall longevity. The physiological roles of DF can
vary depending on nature of the compound. Properties of DF
such as viscosity, molecular weight, molecular size, solubility,
water-holding capacity, and fermentation characteristics by gut
microbiota are known to play a role in the physiological effects
of DF-based functional foods. Increased intake of soluble fibers
is known to suppress hyperglycemia and to improve insulin
sensitivity in nondiabetic and diabetic individuals. Cardiovas-
cular protective roles of DF are imparted by attenuation of
blood cholesterol levels specifically by interfering and limiting
cholesterol fractions transported by LDL. The viscous charac-
teristics of fiber are implicated to have a dominant role in this
regard. Energy load reduction and weight control are imparted
by (1) displacing available energy and nutrients from the diet;
(2) increasing chewing, limiting intake, and promoting
secretion of saliva and gastric juice resulting in an expansion
of the stomach and increased satiety; and (3) reducing absorp-
tion efficiency of the small intestine. Diets with high DF con-
tents are known to have slow nutrient digestion and gastric
emptying rates. This action leads toward attenuation of blood
glucose and helps in the control of type 2 diabetes mellitus.
Some DFs are fermented (prebiotic) completely or partially by
the symbiotic microorganisms of the large intestine to SCFA-
like butyrate, which is the preferred energy source for mucosa
cells. Inulin and oligofructans are not used by potential path-
ogenic bacteria but are the preferred substrate for beneficial
Bifidobacteria and thus help to maintain and restore the balance
of healthful gut microbiota. Beta-glucans and the presence of
magnesium in whole grains and/or bran-rich DF are known to
lead to improved insulin sensitivity to suppress type 2 diabetes
and other metabolic syndrome diseases. Psyllium (or
ispaghula; Plantago ispaghula or Plantago ovata) fibers and oat
and barley b-glucans are used in functional foods targeted at
lower cholesterol and blood sugars for improved cardiovascu-
lar health, control of type 2 diabetes, and weight reduction and
maintenance. Cereal grain DFs are generally recognized to be
more potent than fruits and vegetable fibers in the many
beneficial physiological roles of the DF functional foods.
Excessive DF intake through functional foods or other regular
diets may lead to deleterious effects because it contributes to
reduced absorption of some vitamins, minerals, proteins, and
energy. It may also lead to diarrhea, flatulence, bloating, and
abdominal discomfort. Children less than 1 year old and older
people, in particular, are advised not to consume higher than
recommended DF levels. The recommended AI for total DF is
1925 g day 1 for children aged 18 years, 3138 g day 1 for
boys aged 918 years, 3830 g day 1 for males greater than 19
years old, 26 g day 1 for girls aged 913 years, 2521 g day 1
for females greater than 19 years old, 28 g day 1 at pregnancy,
and 29 g day 1 during lactation.
Whole Grains
Whole grains are defined by AACCI in 1999 as consisting of
the intact, ground, cracked or flaked caryopsis, whose principal
anatomical components the starchy endosperm, germ, and
bran are present in the same relative proportions as they exist
in the intact caryopsis. Thus, whole grains are good sources of
various nutrients, starch, proteins, lipids, DF, vitamins (thia-
mine, riboflavin, niacin, pantothenic acid, vitamin B6, folate,
and vitamin E), minerals, and phytochemicals such as various
phenolic compounds, phytosterols, tocols, lignans, and carot-
enoids. AI of whole grain foods has been associated with GIH
and reduced risk of various metabolic syndrome-related dis-
eases (obesity, diabetes, hypertension, and dyslipidemia) and
various cancers. For example, consumption of two to three
servings per day (about 48 g) of whole grains is preventive of
the risks of CVD, type 2 diabetes mellitus, overweight, and
obesity. Whole grain intake results in greater intake of diverse
nutrients. Although whole grain has been defined, it remains
challenging for researchers, industry, regulatory authorities,
and consumers to clearly identify what constitutes whole
grain foods. Following a review of evidence from various epi-
demiological, interventions, animals, and human clinical stud-
ies on relationship of whole grain intake and the many benefits
 FOOD GRAINS AND WELL-BEING | Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics 13

on health and wellness, an expert panel of whole grain working
committee in 2014 has recommended that 8 g of whole grain
per 30 g serving (27 g per 100 g), without a fiber requirement,
be considered a minimum content of whole grains that is
nutritionally meaningful and that a food providing at least
8 g of whole grains per 30 g serving be defined as a whole-
grain food. This definition also reconciles with the one offered
by AACCI that whole grain food product must contain 8 g or
more of whole grain per 30 g of product. Sources of whole
grain functional foods include whole wheat and rye flours,
barley and whole barley flour, whole grain maize meal,
grits and popcorn, teff, sorghum, oat groats, whole grain
millets, brown rice, quinoa, amaranth, buckwheat, and wild
rice. Thus, whole grain foods (whole grain snacks and
extruded, fermented, and baked foods) can be regarded as
functional foods rich in DF and phytochemicals. Many African
traditional diets and other indigenous foods around the globe
are based on whole grains being consumed as boiled, cooked,
fermented, and baked as in injera from grain teff in Ethiopia.
Such diets can be described as functional foods and they have
been serving communities since folklore.
Probiotics, Prebiotics, and Synbiotics
Human organs particularly the gastrointestinal tract (GIT) are
known to be host for vast numbers of microbes ( 1015 bacte-
rial cells), which are designated as microbiota, microflora, or
normal flora. These microbiota are today extensively exploited
for GIT health and beyond under probiotic and synbiotic
functional foods. Consumption of such diets is actually not
new because a range of naturally occurring prebiotics
and probiotics primarily from the genera Lactobacillus,
Bifidobacterium, and Saccharomyces are consumed throughout
the world in the indigenous fermented foods since folklore.
Probiotics and prebiotics are also available commercially in the
form of dietary supplements and clinical therapeutics to be
delivered orally or nonorally.
Probiotics
The word probiotics is derived from the Greek term pro bios
meaning for life. The widely accepted definition of probiotics
given by FAO/WHO (2001) as Live microorganisms which
when administered in adequate amounts confer a health ben-
efits on the host was recommended in August 2014 by Inter-
national Scientific Association for Probiotics and Prebiotics
(ISAPP) to be used with minor rewording as Live microorgan-
isms that, when administered in adequate amounts, confer a
health benefit on the host. Major bacterial strains commonly
used in probiotics are lactic acid bacteria (LAB) from the genera
of Lactobacillus and Bifidobacterium (Table 1). Other LAB
reported are from Pediococcus, Lactococcus, Leuconostoc,
Streptococcus, and Weissella. Certain yeast and fungal strains
are also recognized for probiotic effects. The ISAPP restricted
the term probiotic to be used only on products that deliver live,
safe microorganisms with suitable viable counts ( 106 colony-
forming units per gram of a product) of well-defined strains
with a reasonable expectation of delivering benefits for well-
being of the host. Ability to survive, proliferate, and colonize
the gut is also a criterion although some cultures lack such
ability like traditional yogurt starter cultures (S. salivarius
subsp. thermophilus and L. delbrueckii subsp. bulgaricus), which
are recognized as probiotics because of their production of
b-galactosidase that helps lactose digestion in maldigesters.
ISAPP has divided live cultures into two: (1) probiotic (non-
oral probiotic foods, probiotic drugs, probiotic medical foods,
nonoral probiotics, defined microbial consortia, probiotic die-
tary supplement, probiotic infant formula, and probiotic ani-
mal feed) and (2) not probiotic (fermented foods with
undefined microbial content and undefined consortia includ-
ing fecal microbiota transplant).
The various beneficial functions of probiotics are (1)
structural and histological such as the development of healthy
structure and morphology of the gut; (2) metabolic such as in
the synthesis of B vitamins, vitamin K, amino acids, SCFA,
antioxidants, and bacteriocins and in the biotransformation of
bile, which help to modulate glucose and cholesterol
homeostasis; and (3) protective from colonization by pathogens
such as through production of antimicrobial compounds, by
competing for nutrients and attachment sites on gut lining. The
various health benefits documented for probiotic diets include
suppression of (1) obesity and type 2 diabetes through modu-
lating glucose metabolism; (2) allergic inflammation and atopic
dermatitis; (3) liver disease; (4) hypercholesterolemia and CVD

Table 1 Microbial species used as probiotics

Species Strain

Lactobacilli L. acidophilus, L. bulgaricus, L. casei, L. delbrueckii ssp. bulgaricus, L. fermentum, L. johnsonii, L. crispatus, L. salivarius,
L. bifidus, L. reuteri, L. plantarum, L. helveticus, L. casei subsp. rhamnosus, L. gallinarum, L. brevis, L. gasseri,
L. cellobiosus, L. vitulinus, L. collinoides, L. cremoris, L. ruminis, L. dextranicum, L. lactis, L. rhamnosus, L. curvatus, L. faecium,
L. paracasei
Bifidobacteria B. bifidum, B. adolescentis, B. brevis, B. longum, B. animalis, B. infantis, B. thermophilum, B. breve, B. essencis, and B. lactis
Bacillus B. coagulans (GanedenBC30), B. lactis DR10, B. licheniformis, B. subtilis R0179, B. subtilis (natto) OUV23481
Pediococcus P. acidilactici, P. pentosaceus, P. halophilus
Lactococcus L. lactis subspp. lactis and cremoris
Leuconostoc L. mesenteroides subsp. dextranium, paramesenteroides, or lactis
Streptococcus S. diacetilactis, S. cremoris, S. lactis, salivarius subsp. thermophilus, S. faecium, S. equinus
Weissella W. cibaria and confusa
Yeast Saccharomyces cerevisiae, S. cerevisiae var. boulardii
Fungi Aspergillus oryzae, Scytalidium acidophilum
14 FOOD GRAINS AND WELL-BEING | Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics
through modulating cholesterol, LDL-cholesterol, and
triglyceride metabolisms; (5) various cancers such as colon,
bladder, and cervical cancers; (6) inflammatory bowel disease;
(7) diarrhea; (8) lactose intolerance; (9) renal diseases; (10)
hormonal immune response failure; and (11) ingestion of
toxic compounds. Although probiotics containing dairy prod-
ucts are dominant, probiotic-fermented cereals and soy bever-
ages, flakes, baked goods, soups, and traditional fermented
cereals are also available. Microencapsulation of probiotics is
currently being researched to enable incorporation into various
grain products. Fermented grain products containing probiotics
have the following advantages: the possibility of grain compo-
nents to be prebiotic sources, multimedia characteristics of the
grain products, and their lactose- and cholesterol-free natures.
Prebiotics
The current consensus definition given is A dietary prebiotic is a
selectively fermented ingredient that results in specific changes,
in the composition and/or activity of the gastrointestinal micro-
biota, thus conferring benefit(s) upon host health. Compounds
recognized as prebiotics include inulin-type fructans (inulin
fructooligosaccharides, and oligofructose), lactulose, galactooli-
gosaccharides, isomaltooligosaccharides, lactosucrose, xylooli-
gosaccharides, glucooligosaccharides, sugar alcohols, resistant
starches, and modified starches. Resistant starches, fructans, ara-
binoxylans, arabinoxylan oligosaccharides, and b-glucans are
reported in various grains such as in wheat, rye, maize, barley,
oat, and maize.
Prebiotic food products are available in the market as bev-
erages (soy milk, smoothies, and juice), bakery products
(snack bars, biscuits/cookies, waffles, and pancakes), breakfast
cereal (extruded cereals and instant oatmeal), confectionery
(chocolate and gummy candy), dairy products (yogurt, ice
cream, and frozen yogurt), desserts (pudding and jelly/jello),
fruit products (fruit preparations), and infant and toddler
foods and in specialty liquid nutrient supplements. Inulin-
type fructans are popular prebiotics. Even though fructans are
present in grains, good sources are from Jerusalem artichoke,
chicory, garlic, and onion.
Prebiotics are fermented to SCFA acetate, propionate, and
butyrate. Other bacterial end products include lactate, ethanol,
succinate, formate, valerate, caproate, isobutyrate, 2-methyl
butyrate, and isovalerate. Bacterial fermentation takes place
in the cecum and colon, where SCFAs are absorbed, stimulat-
ing absorption of vitamins, water, and minerals (calcium and
magnesium). The SCFAs have been known to have a protective
effect on the intestinal epithelium. The colonic bacteria prefer
butyrate as their sole source of energy and are also metabolized
by the colonic epithelium where it regulates cell growth and
differentiation. Butyrate is known to induce secretion of
mucin, antimicrobial peptides, and other factors, which rein-
forces the defense barrier in the colon. Butyrate limitation
leads to inflammation characteristics of the ulcerative colitis.
Acetate and propionate are found in portal blood and are
eventually metabolized by the liver (propionate) or peripheral
tissues, particularly muscle (acetate). Acetate and propionate
might also play a role as modulators of glucose metabolism.
Absorption of such SCFAs is implicated to result in low glyce-
mic responses and to ameliorate insulin sensitivity.
Synbiotics are a combination of probiotics and prebiotics
when administered together. There is synergy between probio-
tics and prebiotics resulting in more effectiveness than using
either one because it favors competitive advantage for probiotic
rapid adaptation in the GIT. Synbiotics are currently available
as combinations of bifidobacteria and fructo-oligosaccharides
(FOS), lactobacillus GG and inulins; bifidobacteria, lactobacilli
and FOS or inulins.
Functional Foods for Lactose Intolerance
Lactose intolerance is the inability to utilize lactose from milk
because of limited lactase enzyme production in the small
intestine in some individuals. Infants can produce adequate
lactase enabling them to utilize lactose of milk, which is the
sole carbon source for the newborn. The ability to generate
lactase enzymes diminishes with age. For individuals with
limited lactase production, lactose will be transferred to the
large intestine intact where it is hydrolyzed and fermented. The
hydrolyzed products cause secretion of water into the intestinal
lumen to balance the osmolarity pressures. This accumulation
of water produces dehydration and electrolyte imbalance on
the systemic side and watery stools on the intraintestinal sides.
The fermentation of the sugar adds gaseous components (car-
bon dioxide, hydrogen, and methane) leading to bloating,
cramping, and flatulence. The limited calcium supply through
inadequate milk intake may lead to osteoporosis. Probiotics
containing Lactobacillus strains are known to recover the lactose
tolerance. The various nondairy probiotic functional foods and
drinks developed include the following:
(1) Soy-based (nonfermented soy-based frozen desserts, fer-
mented soymilk drink, and soy-based stirred yogurt-like
drinks).
(2) Cereal-based puddings; rice-based yogurt; oat-based drink;
oat-based products; yosa, an oat-bran pudding; mahewu, a
fermented maize beverage; maize-based beverage;
probiotic beverages fermented from wheat, rye, millet,
maize, teff, sorghum, and other cereals; malt-based drinks;
boza fermented from cereals, maize, and sorghum; millet
malt-fermented probiotic beverages; and millet or sor-
ghum flour-fermented probiotic beverage.
(3) Fruit- and vegetable-based drinks; fermented banana pulp
and fermented banana juice; beet-based drink; tomato-
based drink; many dried fruits; green coconut water; pea-
nut milk; cranberry, pineapple, and orange juices; ginger
juice; grape and passion fruit juices; cabbage juice; carrot
juice; noni juice; onion puree; probiotic banana puree;
nonfermented fruit juice beverages; and blackcurrant juice.
(4) Others (starch-saccharified probiotic drink, probiotic
cassava-flour product, meat products, and dosa (rice and
Bengal gram)).
Conclusions
Various DFs are available from whole grains, which are also
sources of other nutrients and phytochemicals. The health-
promoting effects of DF (GIH, cancer prevention, and suppres-
sion of metabolic syndrome-related diseases) are widely
acknowledged. Functional foods containing DF are available
 FOOD GRAINS AND WELL-BEING | Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics
as baked, extruded, fermented foods and beverages. Many
components of DF have prebiotic effects. The health-
promoting effects of prebiotics are known by their selective
growth promotion of probiotics, fermentation to SCFA, pre-
vention of various cancers, and enhancement of mineral
absorption. They also suppress CVD, obesity, and diabetes.
Prebiotic functional foods are available as beverages, whole
grains, fermented cereal foods, bakery products, and RTE,
infant, and toddler foods and in specialty liquid nutrient sup-
plements. Probiotics are known to have various healthful
effects (enhancement of gut immunity, anti-inflammation,
prevention of pathogenic infection, and suppression of irrita-
ble bowel syndrome (IBS), cancers, CVD, diabetes, rheumatoid
arthritis, liver diseases, lactose intolerance, and diarrhea).
Currently, nondairy probiotic functional foods and drinks
such as soy-based foods, fermented cereals, and extruded and
baked foods are on the rise. Microencapsulation is one option
to incorporate probiotics in grain products. A combination of
prebiotics and probiotics as synbiotics has synergetic health-
promoting effects. Many traditional fermented grain foods
with live or active cultures if fully characterized have huge
potential to be used as probiotic functional diets. Develop-
ment of grains with desirable DF profiles of prebiotic potential
and improved phytonutrients can advance prebiotic, probiotic,
and DF functional foods. The bioactives used in DF, whole
grains, and prebiotic functional foods are concentrated in the
outer portions of the grains, which may be contaminated by
toxicants that can pose food safety challenges. Good agricul-
tural practices, good manufacturing practices, hazard analysis
critical control points, and adequate grain drying, cleaning,
scouring, and/or removal of very small portion of outermost
layers of grains during milling can suppress such food safety
challenges and also preserve functional ingredients.
Exercises for Revision
Provide definitions of DFs, whole grain foods, prebiotics,
probiotics, and synbiotics.
List up to seven different compounds regarded to be DFs.
List up to four different compounds regarded to be
prebiotics.
Provide genera of microbial species widely used as
probiotics.
Provide essential characteristics of probiotics.
Provide at least two health beneficial effects of DFs, whole
grain foods, prebiotics, probiotics, and synbiotics.
List up to four indigenous fermented foods consumed in
your communities believed to have the effects of prebiotics,
probiotics, and synbiotics.
List potential safe diets for lactose-intolerant individuals.
Exercises for Readers to Explore the Topic Further
What innovations in grain-processing technology can you
envisage to preserve grain bioactive ingredients used to
deliver optimum DFs, prebiotics, and probiotics?
15
How would you provide further characterization of specific
DF components from grains for their health benefits and
prebiotic effects?
Suggest further improvements that may be made to tradi-
tional fermented grain foods to promote their probiotic
value.
Explore possibilities for developing grains using conventional
breeding or through genetic manipulation to provide desirable
DF profiles for prebiotic potential, like inulin-type fructans and
b-glucans.
See also: Beverages from Grains: Fermentation: Foods and
Nonalcoholic Beverages; Bioactives and Toxins: Bioactives:
Antioxidants; Bioactive Compounds in Wheat Bran; Carbohydrates:
Cereals: Chemistry and Physicochemistry of Non-starchy
Polysaccharides; Glycemic Index, Glycemic Load, and Their Health
Benefits; Health Effects of b-Glucans Found in Cereals; Resistant
Starch and Health; Food Grains and the Consumer: Cultural
Differences in Processing and Consumption; Fortification of Grain-
Based Foods; Genetically Modified Grains and the Consumer; Grains
and Health: Misinformation and Misconceptions; Grains and Health;
Labelling of Grain-Based Foods; Food Grains and Well-being:
Functional Foods: Overview; Nutrition: Soy-Based Foods; Food
Grains: Intolerance, Allergy and Diseases: Celiac Disease;
Non-wheat Foods: Soybean: Soy-Based Fermented Foods; The
Basics: Grain: Morphology of Internal Structure.
Further Reading
AACCI Dietary Fiber Definition Committee (2001) The definition of dietary fiber. Cereal
Foods World 46(3): 112126.
Anderson JW, Baird P, Davis Jr. RH Jr., Ferreri S, Knudtson M, Koraym A, Waters V,
and Williams CL (2009) Lead article: Health benefits of dietary fiber. Nutr. Rev.
67(4): 188205.
Binns N (2013) Probiotics, Prebiotics and the Gut Microbiota. Belgium: ILSI Europe.
Birkett AM and Cho SS (2013) Cereal fiber and health: Current knowledge. Cereal
Foods World 58(6): 309313.
Cho SS and Finocchiaro ET (2010) Handbook of Prebiotics and Probiotics Ingredients
Health Benefits and Food Applications. Boca Raton, FL: CRC Press, Taylor &
Francis Group.
Corona-Hernandez R, Alvarez-Parrilla E, Lizardi-Mendoza J, Islas-Rubio AR, de la
Rosa LA, and Wall-Medrano A (2013) Structural stability and viability of
microencapsulated probiotic bacteria: A review. Comprehen. Rev. Food Sci. Food
Saf. 12: 614628.
Dwivedi S, Sahrawat K, Puppala N, and Ortiz R (2014) Plant prebiotics and human
health: Biotechnology to breed prebiotic-rich nutritious food crops. Electron. J.
Biotechnol. 17: 238245.
Ferruzzi MG, Jonnalagadda SS, Liu S, Marquart L, McKeown N, Reicks M,
Riccardi G, Seal C, Slavin J, Thielecke F, van der Kamp JW, and Webb D
(2014) Developing a standard definition of whole-grain foods for dietary
recommendations: Summary report of a multidisciplinary expert roundtable
discussion. Adv. Nutr. 5: 164176.
Foschia M, Peressini D, Sensidoni A, and Brennan CS (2013) Review the effects of
dietary fibre addition on the quality of common cereal products. J. Cereal Sci.
58: 216227.
Granato D, Branco GF, Nazzaro F, Cruz AG, and Faria JAF (2010) Functional foods and
nondairy probiotic food development: Trends, concepts, and products. Comprehen.
Rev. Food Sci. Food Saf. 9: 292302.
Hill C, Guarner F, Reid G, Gibson GR, Merenstein DJ, Pot B, Morelli L, Canani RB,
Flint HJ, Salminen S, Calder PC, and Sanders ME (2014) Consensus statements:
The International Scientific Association for Probiotics and Prebiotics consensus
16 FOOD GRAINS AND WELL-BEING | Functional Foods: Dietary Fibers, Prebiotics, Probiotics, and Synbiotics

statement on the scope and appropriate use of the term probiotic. New York: Slavin J, Tucker M, Harriman C, and Jonnalagadda SS (2013) Whole grains:
Macmillan, Expert Consensus Document, 11, 506-514. Definition, dietary recommendations, and health benefits. Cereal Foods World
Jones JM (2014) CODEX-aligned dietary fiber definitions help to bridge the fiber gap. 58(4): 191198.
Nutr. J. 13(34): 210.
Marsha AJ, Hill C, Rossa RP, and Cotter PD (2014) Fermented beverages with health-
promoting potential: Past and future perspectives. Trends Food Sci. Technol.
39: 135145. Relevant Websites
Poutanen K, Sozer N, and Valle GD (2014) Review: how can technology help to deliver
more of grain in cereal foods for a healthy diet? J. Cereal Sci. 59: 327336. http://wholegrainscouncil.org/whole-grains-101/existing-standards-for-whole-grains
Saad N, Delattre C, Urdaci M, Schmitter JM, and Bressollier B (2013) An overview of the Whole Grains Council.
last advances in probiotic and prebiotic field. LWT- Food Sci. Technol. 50: 116. http://www.isapp.net/Home ISAPP.
 Nutrition: Soy-Based Foods
AM Fehily, Tinuviel Software, Warrington, UK
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 348354, 2004, Elsevier Ltd.
Topic Highlights
Soy is a widely used protein source in Asian cuisine, and the
source for most texturized vegetable protein and vegetarian
meat substitutes.
Soy isoflavones may have significant effects on cholesterol
(and hence on coronary heart disease) reduction; on cancer
risk; and on reduction in menopausal symptoms.
Soy isoflavones, as phytoestrogens, may have negative
effects if consumed at high levels.
Soy contains antinutritional constituents such as lectins,
goitrogens, and phytic acid.
Learning Objectives
To understand the chemical composition of soybean and its
nutritional and antinutritional effects.
To understand the health consequences of isoflavone
consumption.
To be aware of the dietary significance of soy proteins.
Use of Soy in Foods
Soybeans are perhaps the most versatile of plants, providing the
source for a wide range of ingredients. In the Far East, soybeans
are used to produce foods such as tofu, miso, tempeh, and soy
sauce. In Western countries, soybeans have become widely used
in recent years in the production of a range of very diverse
ingredients, e.g., soy flour, soy protein, soy oil, soy lecithin, and
mono- and diglycerides of fatty acids. These ingredients are
widely used in foods. For example, soy flour and soy protein
are used in foods such as breads, pizza bases, other cereal prod-
ucts, and some meat products. Soy protein is often used to make
vegetarian alternatives to meat products, e.g., textured vegetable
protein (TVP) in vegetarian burgers, vegetarian sausages, etc. Soy
oil is used as a vegetable oil both in domestic cooking and by the
food industry. Soy oil is also used in the manufacture of spread-
able fats. Soy lecithin and mono- and diglycerides of fatty acids
are used as emulsifiers in many foods, e.g., breads, fat spreads,
and ice cream. Vegan alternatives to cows milk, cheese, and
yogurt are produced from soy. Soy is also used to produce infant
formulas for those infants who are intolerant to lactose or cows
milk protein.
Composition
Soybeans contain a wide range of nutrients. They contain some
antinutrients and also other components, e.g., isoflavones,
which may have beneficial effects on health.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00066-8
Nutrients
Soybeans contain protein, fat, carbohydrate, vitamins, min-
erals, and fiber.
Protein
Soybeans are a good source of protein, containing 14 g per
100 g. This is more than the protein content in most other
vegetables and in most other legumes (typically 59 g per
100 g), the exception being peanuts which contain 26 g of
protein per 100 g. In terms of protein quality, soy contains
substantial amounts of most essential amino acids. When
compared with a reference protein (egg), soy protein, like
other legumes, is deficient in sulfur-containing amino acids
(the essential amino acid methionine and the nonessential
amino acid cystine) and has a higher content of lysine.
Combining legumes with cereals provides a meal with a high
protein quality, as cereals have adequate amounts of sulfur-
containing amino acids but are deficient in lysine.
Carbohydrate
Soybeans contain 5 g of carbohydrate per 100 g. This is lower
than that of many other legumes. Of the total carbohydrate
content present, 37% is starch, 41% sugars, and 22% oligosac-
charides. The sugars present are sucrose, fructose, and glucose.
Fat
Soybeans contain 7 g of fat per 100 g. This is more than the fat
content in other legumes, most of which are low in fat. The
exception to this is peanuts, which contain 46 g of fat per
100 g. The fatty acid composition of soy has a high ratio of
unsaturates to saturates. Of the fatty acids, 16% are saturates,
24% are monounsaturates, and 60% are polyunsaturates. The
polyunsaturated fatty acids present are the essential fatty
acids linoleic acid (18:2n6) and a-linolenic acid (18:3n3).
Minerals
Soybeans contain significant amounts of several minerals: cal-
cium, iron, magnesium, potassium, phosphorus, and zinc (see
Table 1). One hundred grams of soybeans provides  12% of
the daily reference nutrient intake (RNI) (UK) for calcium,
20% for iron, 23% for magnesium, 15% for potassium, 45%
for phosphorus, and 13% for zinc (see Figure 1).
Vitamins
Soybeans contain a number of water-soluble vitamins: thia-
min, riboflavin, niacin, vitamin B6, and folate (see Table 1).
One hundred grams of soybeans provides  15% of the daily
RNI (UK) for thiamin, 8% for riboflavin, 21% for niacin, 19%
for vitamin B6, and 27% for folate (see Figure 1). Soybeans
also contain vitamin E: 1 mg per 100 g.
17
18 FOOD GRAINS AND WELL-BEING | Nutrition: Soy-Based Foods

Table 1 Vitamin and mineral composition of soybeans (lectins, goitrogens, and digestive enzyme inhibitors). These
substances are, however, inactivated by appropriate cooking of
Amount per 100 g as consumed the beans or, in the case of goitrogens, counteracted by ade-
Sodium 1 mg quate iodine intake.
Potassium 510 mg
Calcium 83 mg Lectins
Magnesium 63 mg Lectins, also known as haemagglutinins, are present in raw
Phosphorus 250 mg soybeans and in other legumes. They are, however, heat labile
Iron 3.0 mg and are inactivated when the beans are properly cooked. In
Copper 0.32 mg experimental animals these polymeric proteins have been
Zinc 0.9 mg shown to cause damage to red blood cells and intestinal
Manganese 0.7 mg
mucosa, and thereby impaired nutrient utilization and loss of
Selenium 5 mg
body weight. In humans, these compounds can result in nau-
Iodine 2 mg
Vitamin A (retinol equivalents) 1 mg sea, vomiting, diarrhoea, and abdominal pain.
Vitamin D 0 mg
Vitamin E 1.13 mg Goitrogens
Thiamin 0.12 mg Enlargement of the thyroid gland has been shown to occur in
Riboflavin 0.09 mg rats fed soybean meal. The goitrogenic agent in soybeans is
Niacin equivalents 2.7 mg unknown. However, the effect is counteracted by adequate
Vitamin B6 0.23 mg iodine intake.
Vitamin B12 0 mg
Folate 54 mg
Digestive enzyme inhibitors
Pantothenate 0.18 mg
Raw soybeans contain certain proteins (protease inhibitors)
Biotin 25.0 mg
Vitamin C Trace which react with digestive enzymes (trypsin, chymotrypsin or
salivary, and pancreatic a-amylase), thereby interfering with
Data from Royal Society of Chemistry and Ministry of Agriculture Fisheries and Food the digestion of protein and starch. In humans, raw soy or
(1991) McCance and Widdowsons The Composition of Foods, 5th edn. Cambridge, isolated protease inhibitors increase levels of cholecystokinin
UK: Royal Society of Chemistry. Crown copyright is reproduced with the permission of (CCK) and pancreatic secretion. It is thought that chronic
the Controller of Her Majestys Stationery Office. pancreatic stimulation may lead to pancreatic hypertrophy,
hyperplasia, and possibly to cancer. In rats, raw soy has been
Folate shown to increase risk of pancreatic cancer. However, protease
inhibitors present in raw soy are inactivated by heat and are
Vitamin B6
therefore not a problem in cooked beans. For example, in
Niacin
countries such as Japan where soy foods are widely consumed,
Riboflavin
the incidence of pancreatic cancer is similar to or less than that
Thiamin for Western countries where soy forms a relatively small com-
Zinc ponent of the total diet.
Phosphorus
Potassium Phytic acid
Magnesium Phytic acid is present in soybeans and also in other fiber-
Iron containing foods. It has been suggested to reduce the absorp-
Calcium tion of calcium, iron, zinc, and vitamin D from the diet.
However, these effects have been observed in in vitro experi-
0 10 20 30 40 50 ments and not in in vivo studies, even at fiber intakes at the
Figure 1 Vitamin and mineral composition of soybeans in relation to upper limit of the normal human consumption range. In
reference nutrient intakes (% of reference nutrient intake (RNI) provided addition, levels of micronutrients tend to be higher in fiber-
by 100 g). (Reproduced from (2003) Encyclopedia of Food Sciences rich foods than in fiber-poor foods. Thus, adverse effects of
and Nutrition, 2nd edn, p. 5393. Elsevier Ltd.) phytic acid on nutrient absorption are likely to be unimportant
at the levels of fiber normally consumed in the human diet.
Fiber
Soybeans are a valuable source of fiber in the diet: 6 g of non- Isoflavones
starch polysaccharides (NSP) per 100 g. Of this, 44% is soluble
fiber and 56% insoluble fiber. One hundred grams of soybeans Soybeans contain isoflavones, a group of compounds that are
provides 33% of the daily dietary reference value (UK) for fiber. structurally similar to oestradiol (phyto-oestrogens). Soybeans
contain higher amounts of the isoflavones genistein and
daidzein (see Figure 2) and smaller amounts of glycitein,
primarily in the form of glycosides: genistin, daidzin, and
Antinutrients
glycitin (total isoflavone content of 180 mg per 100 g). The
Raw soybeans contain a number of substances which may have glycosides are hydrolyzed in the gut by a bacterial enzyme,
detrimental effects on digestion and other metabolic processes glucose being removed to produce the aglycone forms
 FOOD GRAINS AND WELL-BEING | Nutrition: Soy-Based Foods
OH
O balance sheets provide data for the amounts of various foods
HO Daidzein
O
OH
OH O Soybeans can be an important component of a normal
HO Genistein
O For vegetarians, and particularly for vegans who eat no
Figure 2 Chemical structure of the main isoflavones present in soy
daidzein and genistein. (Reproduced from (2003) Encyclopedia of Food
Sciences and Nutrition, 2nd edn, p. 5394. Elsevier Ltd.)
(genistein, daidzein, and glycitein). Daidzein and genistein
have been shown to have weak oestrogenic activity and are
able to bind with low affinity to oestrogen receptors, the affin-
ity being greater for b-receptors than for a-receptors. Compared
with 17b-oestradiol (the main oestrogen produced by the
body), daidzein and genistein have been reported to have
much less potency in producing oestrogenic effects, by about
1000-fold or more.
In the 1940s, it was reported that phyto-oestrogens may
exert adverse effects on uterine and ovarian function. This was
because sheep grazing on pastures, containing a particular type
of clover (Trifolium sp.) rich in formononetin, which is con-
verted to daidzein in the rumen, developed a widespread infer-
tility. An infertility syndrome has also been described in
captive cheetah, as a result of soybean use, the syndrome
being reversed by removal of soy from the food. No adverse
effects of soybeans on human fertility have been reported.
More recently, it has been recognized that isoflavones may
act either as weak oestrogens or as antioestrogens, competing
for oestradiol at the receptor complex, yet failing to stimulate a
full oestrogenic response after binding to the nucleus. This fact
has stimulated much research into whether isoflavones may
have a protective role in hormone-related diseases such as
breast cancer.
Role of Soy in the Diet
Soy has been a staple in the diet of many Far Eastern coun-
tries for centuries, e.g., foods such as tofu, tempeh, and miso.
These traditional soy foods are not very widely consumed in
Western countries, but are sometimes used as an alternative
to meat. As soybeans are extremely versatile, as described
earlier, consumption of foods containing soy has increased
in Western countries over the last 50 years. Nevertheless, the
amount of soy consumed in Western countries such as the
USA and UK is much less than in countries such as Japan and
19
China. The Food and Agriculture Organisation (FAO) food
available for human consumption in different countries.
These show that the amount of soybeans available for use
in foods for humans is 8.5 kg per person per year in China
and 7.9 kg per person per year in Japan. This compares with
less than 1 kg per person per year in countries such as the
USA and UK. In fact, of the soybeans grown worldwide,
only  10% of the total is used in foods for humans, the
majority is used as feed for animals.
healthy diet as they are low in saturates, provide a source of
fiber, and contain a wide range of other nutrients. Soybeans are
also the main source of phyto-oestrogens in the diet. In addi-
tion, soybeans may have a number of potential health benefits
and these are discussed in the following sections.
animal products, soybeans play a very important role in the
diet. They provide a valuable source of protein that can be of
high quality if soybeans are combined with cereals, since the
amino acid compositions are complementary. They also pro-
vide an important source of iron. Low iron intake can lead to
iron-deficiency anemia. Ensuring adequate iron status can be
difficult for vegetarians as, in general, iron from vegetable
sources is less well absorbed than that from animal sources.
However, soybeans provide a source of iron that can be
absorbed as well as that in meat.
For infants intolerant to cows milk protein or to lactose,
soy-based infant formulas can be an invaluable alternative to
cows milk-based formulas. For older children and adults who
are allergic to cows milk or intolerant to lactose, drinks and
other foods made from soy can be very important components
of the diet, helping to ensure that nutrient intakes are adequate
for the maintenance of health.
In common with other legumes, soybeans are also of agro-
nomic importance, as they increase the nitrogen content of the
soil. Although green plants cannot utilize nitrogen in the
atmosphere, there are several species of bacteria, fungi, and
blue-green algae that are able to transform nitrogen in the air
into a form that can be used by plants. An important genus of
nitrogen-fixing bacteria is Rhizobium, which forms nodules in
the roots of legumes. These bacteria live symbiotically with the
legumes, the bacteria obtaining food from the green plant and
the legumes obtaining abundant usable nitrogen compounds
from the bacteria.
Soy and Coronary Heart Disease
Coronary heart disease (CHD) is a major cause of death in
many countries. For example, in the UK, CHD accounts for
 30% of male deaths and 23% of female deaths.
The level of cholesterol in the blood is a major risk factor for
CHD. Soy protein substituted for animal protein in the diet
results in reductions in total plasma cholesterol (by 9% on
average), low-density lipoprotein cholesterol (LDL-C) (by
13% on average), and triacylglycerols (by 11% on average).
High-density lipoprotein cholesterol (HDL-C) is unchanged or
may be slightly increased. The amount of reduction in blood
20 FOOD GRAINS AND WELL-BEING | Nutrition: Soy-Based Foods
lipids tends to be greater among those with the highest plasma
cholesterol levels at baseline.
A great deal of research has been done to identify the
component(s) of soy protein that is responsible for its effect
on blood lipids. Studies using a mixture of amino acids that
duplicate the amino acid profile of soy protein have found
that this does not have the same effect on blood lipids as
the intact protein. However, studies of soy protein with iso-
flavones intact have shown that this has a lipid lowering
effect, whereas soy protein with isoflavones removed has
no significant effect. It has therefore been suggested that
the isoflavones present in soy protein are largely responsible
for the lipid lowering effect of soy protein. The benefits of
isoflavones present in soy protein on cholesterol lowering
have been suggested to be mediated through upregulation
of LDL-receptor activity. Nevertheless, studies of isolated
and purified isoflavones have failed to show that these have
a lipid lowering effect.
Soy may reduce risk of CHD through several mechanisms in
addition to its ability to lower blood lipids (see Figure 3). For
example, soy isoflavones are known to act as antioxidants and
have been suggested to reduce oxidative damage to LDL-C.
A decrease in oxidized LDL particles, which are considered
atherogenic, may reduce the risk of atherosclerosis.
A study of young cynomolgus monkeys showed that the
size of atherosclerotic lesions was 70% less in those fed on a
diet containing soy protein with isoflavones present compared
with those fed on a diet containing soy protein with isofla-
vones removed. This suggests that isoflavones may inhibit
atherogenesis. In another study of surgically postmenopausal
cynomolgus monkeys, soy protein with isoflavones intact was
found to reduce the progression of atherosclerosis. The magni-
tude of the effect was reported to be comparable to that of
postmenopausal oestrogen therapy.
Soybean isoflavones, in particular genistein, impact benefi-
cially on vascular reactivity. Genistein has also been shown,
in vitro, to have a number of other effects whereby the
Reduce oxidative damage to
LDL-C
Inhibit migration &
proliferation of smooth
muscle cells
Beneficial effects on
vascular reactivity
Delay progression of atherosclerosis
Figure 3 Mechanisms through which soy could reduce risk of coronary heart disease (CHD).
development of atherogenesis may be delayed: inhibition of
the migration and proliferation of smooth muscle cells; inhi-
bition of platelet activation and aggregation; and reduction in
platelet serotonin uptake.
In 1999, in response to the available evidence on the ben-
efits of soy protein, the USA Food and Drug Administration
(FDA) announced that it would allow food manufacturers to
label products containing 6.25 g of soy protein per serving as
helping to reduce risk of heart disease, as part of a balanced diet
low in fat and saturates.
Soy and Cancer
Cancer is a major cause of morbidity and mortality. For exam-
ple, in the UK about one in three people will develop cancer at
some time during their life and cancer accounts for about one
in four of all deaths. There are marked differences in death rates
from several types of cancer between Asian countries and many
Western countries. The differences are more striking for
hormone-dependent cancers of the prostate, breast, and
colon/rectum. For example, the risk of dying from prostate
cancer among Japanese men is only one-fifth that of men in
the USA and Japanese women have a breast cancer mortality
rate that is only a quarter of that of women in the USA.
Countries where mortality from prostate and breast cancers
is low have considerably higher intakes of soy than those
countries where mortality from these cancers is high. In animal
models, studies investigating the effects of soybeans on pros-
tate cancer or breast cancer have shown reduced tumorigenesis.
In humans however, evidence is inconclusive. Epidemiological
studies of prostate cancer conducted in Japan and in Japanese
migrants to Hawaii have shown no significant effect of soy
consumption. However, a recent study in the USA reported
that men who drank one cup of soy milk per day had a risk
of developing prostate cancer that was 70% lower than that of
controls. For breast cancer, although some studies have shown
Soy isoflavones Upregulate LDL
receptor activity
Inhibit platelet activation &
aggregation
Reduce platelet
serotonin uptake
Reduce blood lipids
Reduce CHD risk
 FOOD GRAINS AND WELL-BEING | Nutrition: Soy-Based Foods
that soy can help prevent breast cancer, others have found no
significant effect. An increase in menstrual cycle length has
been observed in some studies, in response to soybean con-
sumption, although no change was observed in others. Such an
effect is of potential relevance to the hypothesis that soy may
be protective against breast cancer, because some data indicate
that longer cycles, which are typical of Asian women, are
associated with a reduced risk of breast cancer.
For colo-rectal cancer data are also inconclusive: some stud-
ies showed no significant effect, some reported a protective effect
of soy, and others observed an increased risk with soy consump-
tion. For cancers at other sites, data tend to show that increased
soybean consumption may be protective against lung and stom-
ach cancer. Much more research is needed to confirm whether
soy has a protective effect against cancer in humans.
Isoflavones appear to be the components of soy that would
be most likely to account for any protective action of soy
against cancer, especially in inhibiting the initiation stage of
carcinogenesis (see Figure 4). Genistein has been shown to
suppress the growth of a wide range of cancer cells. It inhibits
certain enzymes that could affect the onset of cancer or the
growth of tumors: tyrosine protein kinase, mitogen-activated
protein (MAP) kinase, and ribosomal S6 kinase. Genistein also
inhibits the DNA repair enzyme topoisomerase II and acts as
an antioxidant, thus potentially preventing oxidative DNA
damage. It increases in vitro concentrations of transforming
growth factor b, which is thought to inhibit the growth of
cancer cells. In addition, genistein has also been shown to
inhibit angiogenesis, the formation of new blood vessels, an
abnormal event that occurs as part of the growth and expan-
sion of malignant tumors. However, many of these effects have
been shown with very high concentrations of genistein and not
in cells treated with levels likely to be achieved in plasma of
human subjects eating soy foods.
Soy and Bone Health
Osteoporosis is a clinical condition in which there is a reduced
amount of bone per unit volume and an increased susceptibil-
ity to fractures, particularly fractures of the vertebrae, distal
forearm (Colles fracture) and hip. Of these, hip fracture is the
most severe, since patients require a prolonged hospital stay,
there is a high mortality rate (about 20% within six months of
the fracture) and of those who survive may suffer permanent
disability and dependency. Osteoporosis is a major health
Soy isoflavones
Women: increase Inhibit activity of
menstrual cycle length various enzymes
Reduce initiation of cancer & growth of cancer cells
Figure 4 Mechanisms through which soy could reduce risk of cancer.
21
problem in many Western countries, although not in Asian
countries. A number of factors are known to increase risk of
osteoporosis, including insufficient dietary calcium, low phys-
ical activity, and lack of oestrogen.
In women, bone mass reduces at a rapid rate in the first few
years after the menopause, due to the reduction in oestrogen.
Hormone replacement therapy (HRT) is well known to reduce
bone loss in postmenopausal women. However, HRT is not
appropriate for all women. Since soy may have oestrogenic
effects, it has been hypothesized that it may provide an alter-
native to HRT.
Soy protein containing isoflavones has been reported to
reduce bone loss due to oestrogen deficiency in ovariectomized
rats. Soybean isoflavones have also shown a protective effect
on bone loss in ovariectomized rats, suggesting that the bene-
ficial effect of soy protein is due to isoflavones. The effect of
isoflavones on bone loss in rats has been reported to be similar
to that for oestrogen. The bone sparing effect of soy protein
isoflavones has been suggested to be due to a reduction in bone
resorption and/or to an increased osteoblast activity (increas-
ing bone formation).
In humans, most but not all studies indicate that soy pro-
tein containing isoflavones favorably affects bone turnover and
bone mineral density in the lumbar spine of perimenopausal
and postmenopausal women. Treatment with ipriflavone, a
synthetic isoflavone, has also been reported to have bone
conserving effects in postmenopausal women with low bone
mass, confirming that the beneficial effect of soy protein is
likely to be due to isoflavones.
In addition to the effect of soy protein on bone mineral
density, it has also been suggested that soy protein, when
substituted for animal protein, may indirectly enhance bone
strength. Another effect of soy protein is that it helps to con-
serve calcium by reducing urinary calcium excretion. This is
due to the lower sulfur amino acid content of soy protein.
Soy protein therefore appears to have modest beneficial
effects on bone density. However, studies to date have been
short term and have involved only small numbers of subjects.
In addition, no study has investigated whether soy protein
containing isoflavones has an effect on fracture risk.
Soy and Menopausal Symptoms
Hot flushes (also known as hot flashes) are a common symp-
tom among menopausal women in Western societies, but are
Prevent DNA Inhibit
damage angiogenesis
22 FOOD GRAINS AND WELL-BEING | Nutrition: Soy-Based Foods
reported to be much less common in Japan. Whether this
difference is due to the higher consumption of soy in Japan is
not clear. Hormone replacement therapy (HRT) generally alle-
viates hot flushes and other menopausal symptoms such as
vaginitis. There is much research interest in the possibility that
soy may provide an alternative to HRT in this regard.
Evidence of benefit of soy in menopausal women is conflict-
ing. Some researchers have reported an improvement in the
frequency of hot flushes in women taking soy protein daily.
Others have reported no effect of soy protein on the number of
hot flushes experienced but found a reduction in the severity of
symptoms. Others have reported a reduction in both the inci-
dence and severity of hot flushes with soy protein or an isoflavone
extract compared with a control group. Others have reported no
difference in either the frequency or severity of hot flushes in the
intervention group compared with the control group.
Two studies have investigated the effects of phyto-oestrogen
supplements on vaginal cytology and found an increase in cell
proliferation (an indication of oestrogenic activity) and reversal of
menopausal atrophy. Others have reported no significant effect.
There are difficulties in interpreting the results of these
studies. This is because there are differences in the amounts
of soy protein and isoflavones used and differences in the
duration of the studies. Interpretation is also complicated by
the fact that reported frequency and severity of symptoms
tends to reduce in the control group as well as the intervention
group. Thus, much more research needs to be done before soy
protein can be proposed as a potential alternative to HRT for
the control of menopausal symptoms.
Exercises for Revision
Complile a summary of the effects of phytoestrogens on
young adult men and women.
What are the effects of high temperature processing on soy
nutritional characteristics?
Exercises for Readers to Explore the Topic Further
Investigate further the current state of knowledge on the
effect of soy isoflavones on a medical condition of interest
to you.
How would you design a soybean for optimum nutritional
and nutraceutical composition.
See also: Agronomy of Grain Growing: Soybean: Agronomy;
Food Grains and the Consumer: Genetically Modified Grains and
the Consumer; Non-wheat Foods: Soybean: Soy-Based Fermented
Foods; Soybean: Soymilk, Tofu, and Okara; Oilseed and Legume
Processing: Soybean: Processing; Soybean: Soy Concentrates and
Isolates; The Legumes and Pseudocereals: Beans: Overview;
Grain Legumes and Their Dietary Impact: Overview.
Further Reading
Anderson JJB, Anthony M, Messina M, and Garner SC (1999) Effects of
phyto-oestrogens on tissues. Nutrition Research Reviews 12: 75116.
Anderson JW, Johnstone BM, and Cook-Newell ME (1995) Meta-analysis of the effects
of soy protein intake on serum lipids. New England Journal of Medicine
333: 276282.
Bingham SA, Atkinson C, Liggins J, Bluck L, and Coward A (1998) Phyto-oestrogens:
where are we now? British Journal of Nutrition 79: 393406.
Lissin LW and Cooke JP (2000) Phytoestrogens and cardiovascular health. Journal of
American College of Cardiology 35: 14031410.
Messina MJ (1999) Legumes and soybeans: overview of their nutritional profiles and
health effects. American Journal of Clinical Nutrition 70(suppl): 439S450S.
Messina M and Messina V (2000) Soyfoods, soybean isoflavones and bone health: a
brief overview. Journal of Renal Nutrition 10: 6368.
Messina M, Persky V, Setchell KDR, and Barnes S (1994) Soy intake and cancer risk: a
review of the in vitro and in vivo data. Nutrition and Cancer 21: 113131.
Potter SM (2000) Soy new health benefits associated with an ancient food. Nutrition
Today 35: 5360.
Royal Society of Chemistry and Ministry of Agriculture, Fisheries and Food. (1991)
McCance and Widdowsons. The Composition of Foods, 5th edn. Cambridge, UK:
Royal Society of Chemistry.
Relevant Websites
nutrition.org.uk http://www.nutrition.org.uk British Nutrition Foundation.
food.gov.uk http://www.food.gov.uk Food Standards Agriculture Organisation.
apps.fao.org http://apps.fao.org Food and Agriculture Organisation (FAO).
ajcn.org http://www.ajcn.org American Journal of Clinical Nutrition.
 Food Grains and the Consumer

Contents
Grains and Health: Misinformation and Misconceptions
Consumer Trends in Grain Consumption
Cultural Differences in Processing and Consumption
Fortification of Grain-Based Foods
Genetically Modified Grains and the Consumer
Labelling of Grain-Based Foods
Grains and Health

Grains and Health: Misinformation and Misconceptions

DB Sheats and JM Jones, St. Catherine University, St. Paul, MN, USA
2016 Elsevier Ltd. All rights reserved.

Topic Highlights
Claims suggest that wheat is the product of genetic research
and that we are eating genetically altered wheat. In reality,
genetically modified wheat is not sold legally anywhere in
the world.
Claims suggest that wheat and its gluten have changed and
that is why celiac disease, an autoimmune disorder, is on
the rise. While celiac disease rates are increasing, and theo-
ries abound, as to why.
Claims suggest that wheat is addictive. There are no data to
support claims that removing wheat from the diet results in
withdrawal symptoms.
Claims suggest that wheat causes uncontrollable overeat-
ing. Existing data show that ingestion of wheat protein
causes the release of satiety hormones.
Claims suggest that wheat ingestion leads to a host of
mental health issues. There are scanty scientific data to
support these claims.
Claims suggest that wheat causes obesity and diets that
eliminate wheat and other carbohydrates are the most effec-
tive way to lose weight. In reality, any significant weight loss
that occurs with these diets is due to the reduction in total
calories from any source, not just wheat or carbohydrates.
Claims suggest that the elimination of wheat from the diet
results in the cure or improvement of many health condi-
tions. In reality, many of these conditions are made better
by weight loss and not specifically the elimination of wheat.
Claims suggest that the nutrient contribution of wheat and
grains can easily be obtained from other food groups. While it
is true that these nutrients are found in other foods, studies
on nutritional quality indicate that diets without whole and
enriched grains are often lacking in these nutrients.
Food and nutrition professionals need to be able to counter
unfounded theories about wheat and other grains with
sound science and unbiased, critical thinking.
Learning Objectives
To identify current controversies surrounding wheat, grains,
and health.
To describe scientific evidence regarding wheat, grains, and
health.
To understand the important role that food and nutrition
professionals play in dispelling common misinformation
and misconceptions about wheat and grains.
Introduction
Recently, grain consumption, especially wheat, has been
blamed for obesity, diabetes mellitus, other chronic diseases,
food addiction, and impaired brain function. Popular press
books such as Wheat Belly by William Davis, the Paleo Diet by
Loren Cordain, and Grain Brain by David Perlmutter and other
authors and food bloggers claim that eliminating grains is the
key solution to improving health. A review of nutrition and
medical literature suggests a different scenario. This article pro-
vides a review of the scientific evidence regarding grains and
health to dispel misinformation and misconceptions. This
piece will focus on wheat because it has been subject to the
most misinformation.

Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00078-4 23

24 FOOD GRAINS AND THE CONSUMER | Grains and Health: Misinformation and Misconceptions
Wheat Breeding and Genetics
Wheat originated in the Middle East thousands of years ago.
According to the Food and Agriculture Organization (FAO),
wheat is grown on more land area than any other commercial
crop and continues to be the most important food grain source
for humans. Together, rice, wheat, and maize provide over half
of the worlds calories.
Popular press authors and food bloggers suggest that wheat
is the product of genetic research, and today, we are eating
genetically altered wheat. They contend that wheat and its
gluten have changed and our bodies can no longer digest the
modified grains of today, and that is why people are having so
much difficulty digesting wheat and why celiac disease, an
autoimmune disorder, is on the rise.
It is alleged that modern wheat has changed to be less
healthy because of breeding practices. In fact, wheat breeding
is an age-old practice. Significant advances in plant breeding
occurred in the 1940s with the work of Norman Borlaug and
others, using traditional techniques (non-genetically modified
organisms). Lines selected provide increased yield through
disease, drought, and insect resistance, especially in developing
countries. Despite assertions to the contrary, these lines were
produced by crossing wheat varieties with other wheat varieties
from all over the globe. In 1970, Dr Borlaug with his so-called
Green Revolution won the Nobel Peace Prize for his wheat and
grain breeding programs, which produced grains with high
yields that grow under a wide variety of conditions and help
address world food supply challenges.
It is also alleged that genetic modification of wheat has
made it less healthy and has changed the gluten proteins.
This is untrue for two reasons. First, no ill effects can come
from genetically modified wheat because there is none sold
legally anywhere in the world. Second, recent studies from
USDA labs and labs at the University of Saskatchewan and
studies in Europe using grains from seed banks going back as
far as Canadian Red Fife in 1857 show wheat gluten has not
changed.
Concern has been raised about the use of hybrid dwarf
wheats. They now comprise 99% of the wheat grown world-
wide. Short-straw naked (dwarf) wheats are one of the
advances that were part of the green revolution. They increased
yield for two reasons: (1) wheat varieties with long, spindly
straws lodge, meaning that the kernel head is heavy and falls to
the ground, making harvest difficult, and (2) shorter straw
means that inputs, for example, water and nutrients, go into
making wheat kernels rather than straw.
Popular press authors imply that new wheat varieties and
their proteins are unique or harmful in some way. Further, they
posit that 5% of the proteins in wheats are unique, meaning
they are found in neither parent, and that this unexpected
genetic rearrangement results in altered proteins with poten-
tially toxic effects. It is important to consider that plants can
only express proteins that they have the genetic code to pro-
duce. Creating a unique protein requires a mutation of the
DNA or RNA. While environmental conditions can promote
or inhibit the expression of certain proteins, it cannot code for
proteins that are not in the genome. Therefore, hybridization
of wheat does not create unique proteins.
Ancient wheats such as einkorn are held as models of what
we should eat because they have not been hybridized and
because of their protein makeup. In fact, one author says that
einkorn contains 28% protein compared with average protein
contents of 1215% in modern wheats and that ancient wheats
did not cause the symptoms that new varieties do. However,
studies carried out at the University of Saskatchewan where
ancient and other varieties of wheat were grown on adjacent
plots actually show that the crude protein content of einkorn is
18% and that of other ancient and modern varieties had pro-
tein values ranging from 16.3% to 17.5%. This fits well with
the USDA World Wheat Collection, which shows approxi-
mately a threefold variation in protein content of wheat from
7% to 22%, with one-third of this under genetic control and
two-thirds of this controlled by environmental conditions.
Celiac and Related Disorders
Celiac disease is on the rise, as are other autoimmune diseases,
in Western countries. This fact is documented in the scientific
literature. Studies by Mayo Clinic gastroenterologist Joe
Murray and colleagues compared celiac titers in blood samples
from the recent US Air Force recruits with stored blood
samples taken from recruits more than 50 years ago. The
analysis showed 0.2% of recruits had the gene for celiac disease
in 1950 compared with 0.9% of recent recruits, which is a
quadrupling of the incidence rate. Data from Finland also
show an increase from 1% to 2% or a doubling of the rate of
celiac disease incidence in that population. While celiac disease
rates are increasing, it is uncertain whether this is due in part to
better identification and awareness of the disease, as well as a
myriad of other dietary, immunologic, and environmental
changes, especially those that impact the microorganisms and
integrity of the gut. Some theories include the clean theory,
poor diets overall, overuse of antibiotics and other drugs espe-
cially acid-lowering drugs, lack of breast-feeding, and frequent
use of Cesarean sections.
Davis and other popular press authors posit that the reason
for the increase in celiac disease is attributed to the fact that
celiac disease-triggering proteins are expressed at higher levels
in current wheat varieties than found 50 years ago. Such
authors suggest that high-molecular-weight (HMW) glutenins
and higher levels of gliadin of modern wheat are the problem.
While studies show that the immune system reacts to break-
down products of HMW glutenins, data are lacking that show
that these proteins trigger more reactions than those of ancient
wheats or even wheat varieties from 50 years ago. Interestingly,
gliadins are found in higher amounts in ancient versus modern
wheats.
Gluten-containing grains are also implicated in non-celiac
gluten sensitivity (NCGS). This newly recognized disorder has
no medically agreed-upon test to confirm a diagnosis, and
therefore, its existence and incidence are subject to much
debate. The best estimates are that 46% of the population
may have NCGS. NCGS is thought to be present in about 5%
of irritable bowel syndrome cases and may cause the rash,
dermatitis herpetiformis, which can be seen in some celiac
patients.
 FOOD GRAINS AND THE CONSUMER | Grains and Health: Misinformation and Misconceptions
Allergens
Wheat has long been categorized as one of the big eight
allergens (the most common allergens in Western countries).
Despite beliefs that the incidence is much higher, only 0.5% of
children and adults in the United States suffer from wheat
allergy, which can be caused by any number of proteins in
wheat. Glutenins are the most frequent allergens, but gliadins,
especially gamma-gliadin, result in the most severe allergic
reactions. A particular protein, omega-5 gliadin, is responsible
for wheat-dependent, exercise-induced anaphylaxis and may
be the offending protein in the wheat allergies of young chil-
dren. There are also reported allergies to albumins, globulins,
and enzyme inhibitors. Interestingly, delayed introduction of
solid foods including grains such as wheat and rye has been
found to be responsible for greater allergic sensitization in
young children.
Addiction and Overconsumption of Food
Popular press authors and bloggers suggest that wheat is the
worlds most destructive dietary ingredient because during its
digestion, it breaks down into peptides that act as exorphins
(exogenous opioids). These authors assert that wheat is unique
in this role. Studies conducted by the National Institutes of
Health (NIH) in 1979 show that digestion of wheat proteins
can produce peptides that interact with opioid receptors.
However, the claim that wheat is unique in this regard is
incorrect, as these same studies show that other food proteins
also produce peptides that interact with opioid receptors.
Hydrolysates of milk proteins (alpha-, beta-, or kappa-casein;
alpha-lactalbumin; beta-lactoglobulin; and lactotransferrin)
show the highest opioid activity. In addition to milk proteins
and wheat gluten, rice albumin, bovine serum albumin, and
even a protein from spinach all produce peptide fragments
capable of interacting with opioid receptor ligands. It is impor-
tant to consider that the studies indicating wheats possible
opioid potential were conducted either in vitro or by feeding
the preformed peptides, not by feeding wheat itself. Studies
feeding wheat foods, not hydrolysates, must be conducted to
determine the actual effects of peptides from gluten digestion.
Some studies have also reported beneficial effects of these
peptides. If bioavailable, they have the potential to help con-
trol blood pressure and to improve learning performance.
Another theory promulgated by popular press authors and
bloggers is that wheat opioids are highly addictive causing
individuals to lack control of their eating, with subsequent
removal of wheat from the diet producing withdrawal symp-
toms. The controls over hunger and satiety are complex being
affected by many mechanisms from physical feelings of full-
ness (stomach distention) to neuroendocrine, psychosocial,
and sensory factors. While some suggest specific foods such
as sugars and fats are addictive, supporting evidence is
weak and scarce, with no human data. Similarly, there are no
data to support claims that removing wheat from the diet
results in withdrawal symptoms.
Claims from Davis and others that wheat causes uncontrol-
lable overeating conflict with existing data that show wheat
25
protein ingestion causes the release of satiety hormones chole-
cystokinin and glucagon-like peptide 1. Wheat and pea pro-
teins show a stronger ability than other sources to stimulate the
release of both hormones. Some data suggest that consump-
tion of proteins such as those in gluten may be a good dietary
strategy for weight management.
Mood, Mental Fogginess, Autism, Attention-Deficit/
Hyperactivity Disorder, and Schizophrenia
A common assertion in the popular press is that wheat inges-
tion alters mood and causes mental fogginess. There are little
data showing that wheat consumption alters mood or mental
acuity. In a small study of patients with celiac disease, a gluten-
restricted diet failed to improve neurological symptoms. In
contrast, increased serotonin is associated with a sense of
well-being and elevates mood. In a study of malnourished
Indian primary school children, wheat biscuits added to their
diets actually improved cognitive ability. There are data sug-
gesting adding lysine to grain-based diets may reduce measures
of anxiety. In a study of Syrians with marginal diets based
primarily on wheat, the addition of lysine, wheats limiting
amino acid, reduced symptoms of anxiety. This study supports
the fact that complementary plant proteins are needed in grain-
based diets low in complete proteins.
Wheat is also alleged to cause of autism spectrum disorder,
attention-deficit/hyperactivity disorder (ADHD), and schizo-
phrenia. Case reports and other narratives suggest there may
be a link between autism and celiac disease. However, docu-
mentation in the medical literature is scarce. Regarding autism,
there is only one randomized clinical trial and its findings
showed no significant difference in symptoms due to gluten
removal from the diet. A review considered the gold standard
in the medical literature concluded that further large-scale,
randomized-controlled trials are needed. According to
Fasano, a noted celiac disease researcher at the Center for Celiac
Research in Baltimore, MD, while the gluten-free, casein-free
diet is one of the most popular diets for the treatment of autism,
the efficacy of this diet is inconclusive.
Similarly, studies examining the relationship between
wheat and ADHD are lacking, even though sensitivity to a
range of foods, including wheat, has been suggested in a num-
ber of case reports. Studies with very small numbers of subjects
show no improvement in ADHD symptoms with a gluten-free
diet. Screenings for celiac disease in a clinical report (no
control group) involving 67 people diagnosed with ADHD
showed the incidence to be slightly higher than the population
as a whole with 1 out of 7 (vs. slightly < 1 in 10) having celiac
disease. These celiac disease-positive participants showed a
significant decrease in ADHD symptoms.
Schizophrenias link to gluten was noticed during World
War II when a British physician observed decreased admissions
for this disorder. Dr. Dohan theorized that this decrease corre-
lated with decreased wheat consumption imposed by rationing
and postulated links between wheat, celiac disease, and
schizophrenia. Part of the theory suggests that wheat ingestion
affects tight junctions and reduces the guts capacity to prevent
the entry of exogenous substances, thus allowing the develop-
ment of schizophrenia and other mental conditions. It is also
26 FOOD GRAINS AND THE CONSUMER | Grains and Health: Misinformation and Misconceptions
known that schizophrenia incidence increases in the presence
of any autoimmune disease or a history of severe infections. It
is thought that the antibodies produced can impact the brain.
In some studies, a subset of schizophrenia patients show ele-
vated gliadin antibodies. However, classic celiac disease diag-
nostics, either antibodies or gene markers, were not found in
higher rates in schizophrenia patients than in the public at
large.
A comprehensive review looking at the connection between
gluten and schizophrenia showed that gluten withdrawal
resulted in a drastic reduction or full remission of symptoms
but only among a small subset of schizophrenia sufferers.
Thus, in a small group of schizophrenia patients, removal of
wheat may be helpful but not the miracle cure indicated by
Davis and others.
Obesity and Weight Loss
Obesity rates have increased dramatically around the world
since the 1970s. Obesity rates in the United States have
increased by 214% since 1950. Davis and other popular press
authors note that obesity was rare in the 1950s and cited
increased wheat consumption as a cause. It is true that wheat
consumption increased from 1970 to 2000; however, it
dropped between 2000 and 2014. Importantly, these authors
failed to note that during this same period, portion size and
caloric intake have increased dramatically while physical activ-
ity has decreased and screen time has increased. In fact, in 2008
in the United States, there were on average 600 more calories
available per person than in 1970.
Food bloggers and popular press authors suggest that the
proliferation of wheat products in the diet parallels increased
weight and central obesity. Central obesity, also known as
visceral adipose tissue (VAT), has increased in the population
and is a cause for concern due to its association with higher
rates of diabetes, high blood pressure, heart disease, and cer-
tain forms of cancer. While VAT has increased, it is not true that
wheat causes this condition or that the elimination of wheat
will cure this condition. It is well documented that no one food
or food group is responsible for VAT. Instead, ingesting too
many calories of any kind coupled with too little exercise will
result in VAT. Recent data from the Framingham Heart Study
cohort refute claims that wheat increases VAT. Those who had
the least VAT ate two servings per day of refined grains and
three servings per day of whole grains.
For many years, various low-carbohydrate diets have been
recommended as the most effective way for individuals to lose
weight. It is not surprising that eliminating wheat from the diet
has recently been considered to be the holy grail of weight loss
regimens using low-carbohydrate diets. While it is true that low-
carbohydrate diets have been shown to cause more rapid weight
loss than other diets in the first 6 months, these diets do not result
in greater weight loss over time. In addition, low-carbohydrate
diets often result in more dropouts than other diets that are more
balanced and do not eliminate entire food groups. Any significant
weight loss that occurs with these diets cannot be attributed to the
magic of eliminating grains, but instead, it is due to the reduction
in total calories from any source.
Grain Foods and the Glycemic Index
The glycemic index (GI) is a measure that compares the blood
glucose response produced by 50 g of available carbohydrate
from a food with the blood glucose response produced by 50 g
of glucose. Popular press authors and food bloggers correctly
noted that wheat bread has a GI that is higher than table sugar.
However, GI is most often misunderstood because it is used to
compare very different amounts of food. For example, 50 g of
table sugar (approximately three tablespoons) would yield
50 g of available carbohydrate. In contrast, 50 g of available
carbohydrate from whole wheat bread is much more than 50 g
of bread since bread is not all carbohydrate and not all the
carbohydrate is available. Thus, it would take 144 g of whole
wheat bread (5.1 slices at 28 g per slice) or 111 g of white bread
(3.9 slices) to yield 50 g of available carbohydrate.
Grain Foods and Other Diseases
Popular press authors and bloggers claim that elimination of
wheat from the diet results in the cure or improvement of
many conditions including diabetes, coronary heart disease,
multiple sclerosis, rheumatoid arthritis, asthma, acid reflux,
irritable bowel syndrome, ulcerative colitis, acne, and other
rashes. However, many of these conditions are made better
by weight loss, so attributing improvement to wheat removal
is overly simplistic and is likely an inaccurate deduction. A
number of meta-analyses and reviews show that those who
eat grains and whole grains have lower risk or most chronic
diseases with a recent one showing a reduction in both the risk
of cardiovascular and all-cause mortality.
Nutrient Composition of Wheat and Grains
Many diet authors bashing grains and wheat assert that the
nutrient contribution of wheat and grains can easily be
obtained from other food groups. While it is true that most
of the constituents of grains can be found in other foods, there
are some issues. For example, grains provide many of the B
vitamins. While it is true that these are found in other foods,
studies on nutritional quality indicate that diets without whole
and enriched grains are often lacking in these nutrients. The
concern is especially important for folate. Folate fortification of
grains has decreased neural tube defects and other neurological
problems in North American populations by 50% since forti-
fication of grains was mandated. Other compounds such as
betaine are found in wheat and wheat bran and germ, the
highest sources of this important phytochemical. Betaine is
not widely distributed in the foods supply and has importance
in human nutrition.
Diets without grains can have significant problems getting
adequate dietary fiber and a good mix of fiber types. Wheat
bran/fiber is used as the standard against which government
agencies test the efficacy of a fiber in terms of laxation, as it is
the fiber with significantly greater effect on laxation than fibers
from fruits and vegetables. The same fiber in barley and oats
with proved cholesterol-lowering properties is not widely found
 FOOD GRAINS AND THE CONSUMER | Grains and Health: Misinformation and Misconceptions
in many fruits or vegetables, and when found, its structure is
such that it does not have cholesterol-lowering properties.
Further epidemiological studies show that cereal fiber
reduces the risk of certain gut cancers while fruit and vegetable
fibers do not. These data suggest that balance as prescribed in
USDA MyPlate provides the best diet.
Conclusion
Numerous health problems including obesity, diabetes melli-
tus, other chronic diseases, food addiction, and impaired brain
function have been attributed to grains such as wheat. A review
of nutrition and medical literature reveals that there are little
evidence to support claims that wheat and grains cause adverse
health effects and chronic disease and much evidence to show
that a balanced diet that includes wheat and grains, especially
whole grains, reduces chronic disease risk and lowers overall
mortality.
Popular press authors and food bloggers claim that elimi-
nating wheat grains is essential for optimal health for all. While
it is true that wheat and other gluten-containing grains must be
eliminated for those with reactions to gluten and wheat
because of celiac disease, allergies, or other conditions, it is
not true that wheat and grain elimination is best for all.
Further, it is also true that many diets contain too many calo-
ries and grain-based desserts and snacks contribute to excess
calories, and it is not true that the grains and wheats are the
culprit. Rather, it is the poor food choices that should be
addressed. Data for elimination of grains for all are, at best,
scanty and do not show scientific rigor. Food and nutrition
professionals need to be able to counter unfounded theories
about wheat and other grains with sound science and unbi-
ased, critical thinking. This article provides a review of the
scientific evidence regarding grains and health to dispel com-
mon misinformation and misconceptions.
Exercises for Revision
Take a specific wheat controversy and describe the scientific
evidence to support or refute this claim.
Model a diet with and without grains. Calculate the nutri-
tional value of the model diet using a computer analysis to
show what nutrients might be at risk. Show what foods
would need to be consumed in larger quantities. If this
were done, what would happen to calories?
Calculate the calories for 3 days of diets with wheat or
grains included and the calories of that same diet without
the wheat or grains. Show how much weight would likely
be lost with the elimination of these products.
Go to a store and compare the cost, nutritional value, and
eating quality of products that are labeled as gluten- or wheat-
free with their gluten- or wheat-containing counterparts.
Exercises for Readers to Explore the Topic Further
Why is misinformation around diet so readily accepted?
Show the evidence supporting the use of wheat and grains
in preventing any one of the chronic diseases. Assess this
evidence in and evaluate evidence for their nonuse.
27
Assess the evidence associated with grains, carbohydrates,
and brain function. How does this evidence compare with
other nutritional impacts on the brain?
Use the GI tables and look up the values for wheat and
grain staples such as sourdough and whole grain bread
basmati rice and pasta. Explain why such products often
with the same ingredients have varying GIs. Then look at
the amount eaten for 50 g of carbohydrate and compare
that and the nutritional quality of indulgent and other
foods with lower GIs such as candy or chocolate cake.
Using this research, explain when the GI might be useful
for food selection and when it might not yield optimal
nutrition choices.
See also: Appendix 3: Grains, Foods, and Ingredients Suiting
Gluten-Free Diets for Celiac Disease; Carbohydrates: Glycemic
Index, Glycemic Load, and Their Health Benefits; Food Grains and
the Consumer: Grains and Health; Food Grains: Intolerance,
Allergy and Diseases: Celiac Disease; Cereal Allergens; The Gluten-
Free Diet; Grain Composition and Analysis: The Composition of
Food Grains and Grain-Based Products; The Cereal Grains: Wheat:
An Overview of the Grain That Provides Our Daily Bread.
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28 FOOD GRAINS AND THE CONSUMER | Grains and Health: Misinformation and Misconceptions
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 Consumer Trends in Grain Consumption
JM Jones and DB Sheats, St. Catherine University, St Paul, MN, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Trend drivers in developed countries are different from
those in emerging markets and developing countries, but
each can influence the other.
Grain consumption worldwide is increasing, but direct
consumption is decreasing as wealth increases and animal
foods are selected.
Gluten-free is needed for wheat allergy and celiac disease
and related disorders but is unnecessary and expensive. It
may be lower in nutrients especially dietary fiber and folate
for those who do not need to adopt this diet.
Low-carbohydrate, antigrain, and other fad diets impact
grain consumption.
Underconsumption of whole grains and overconsumption
of grain-based desserts and snacks are trends that need to be
addressed in many regions.
The selection of alternate grains and sprouted grains is
another emerging trend.
Learning Objectives
To identify drivers that affect grain consumption and be
able to show how those are different in different regions
and economies,
To name a number of sources of data, which track grain
production and consumption,
To describe drivers of grain consumption for specific grains,
To distinguish between drivers that are trends and those
that are fads and to attribute causes for both.
Introduction
Taste, cost, availability, convenience, and nutrition have a
strong impact on consumer behavior and consumption with
regard to food and grains. Many new trend drivers have been
added or are exerting more influence than in the past. These
include environmental concerns, concerns about food sustain-
ability, the foodenvironment and foodfuel connections,
food provenance and the locavore movement, social norms
that affect attitudes toward growers and producers of food
including concerns about large agribusiness and spawning
the growth of local markets and consumer-supported agricul-
ture, the role of women in the society, the importance of food
and the time spent devoted to food in a culture, the attitudes
toward maintaining food traditions versus encouraging the use
of new foods and methods of production and cookery, glob-
alization of foods, fear and misunderstanding of technologies
and practices involved in food production, distrust of long
esteemed food staples such as grains, and worries about the
rise in obesity and certain other diseases. All of these must be
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00072-3
balanced with the looming specter of inadequate food supplies
to feed the anticipated population of 2050.
Consumer attitudes and ideas change continually, but these
changes seem to occur at a faster rate in the age of social media,
blogs, battling experts, 24 h news cycles, and talk shows,
all which report the latest research findings sometimes with
varying degrees of accuracy. If these voices persist, the
information regardless of its veracity causes changes in
food and grain consumption.
Perception is reality and consumer beliefs and trends drive
the marketplace. Thus, it is important that those producing and
manufacturing grain in planning for the long term be able to
(1) differentiate between fads and trends, (2) to tailor food
product development and market launches so that product
introduction and supply meet consumer expectations, and
(3) to predict changes in the future that allow innovation and
are ready with supply chain and production to meet these
changed demands.
International Consumer Trends
The trend toward consumption of higher-value food products
across all income levels has risen in many countries in recent
years due to higher disposable incomes. While changes in
income affect consumption patterns, increased urbanization
is an important driver. It often results in higher levels of edu-
cation, in more women working outside the home, and in
shifts from jobs that require much energy expenditure to
more sedentary ones, and it may alter incomes and food
choices both positively and negatively. Those trapped in
urban ghettos may shift their diets to cheap, readily available
calorie sources that may be poor choices in terms of nutrient
density. However, consumers thriving in emerging economies
such as China and Mexico shift their food purchases away from
traditional diets and grains and other carbohydrate-rich staple
foods to more expensive sources of calories, such as meat and
dairy products, more packaged foods such as breakfast cereals
and other frozen, instant or ready-to-eat options, and more
foods prepared away from home. According to Marketsand-
Markets, the most rapid rate of growth in the breakfast cereal
category is expected to occur in the Asia Pacific region because
of some of the aforementioned factors.
Some consumption trends affect health either positively or
negatively. The trend reflecting the demand for healthier
cereals and cereal products is happening in many regions.
However, there is also the trend away from healthy, grain-
based staples to increased consumption of grain-based des-
serts, pastries, and snacks, which may add little in terms of
nutrition and may add many unneeded calories.
In terms of food and grains, urbanization often also means
a mixing of people from various regions and ethnic groups
introducing new foodstuffs and tastes and creating the
29
30 FOOD GRAINS AND THE CONSUMER | Consumer Trends in Grain Consumption
availability of a wide array of food products. Many jobs also
require travel that introduces the population to more tastes,
different grains, and different ways of using grains. Many
multiethnic cities have markets and restaurants that offer an
array of grain products from amaranth cereal, to buckwheat
groats, to corn tortillas, to porridge (oat meal), to quinoa pilaf,
to rye crisp, to sorghum beer, to rice noodles, to breads from all
over the world.
Tracking Consumer Trends
Governments have traditionally tracked the supply and avail-
ability of grain-based foods. This is done in a variety of ways.
Government and international food and agriculture bodies
such as WHO/FAO collect statistics that give information on
types and varieties of grain and other products grown; cost and
availability of seeds and inputs; stores of grain; weather,
environmental, and other factors that affect crop planting,
yield, and market availability; food disappearance and
consumption; health issues; and economic factors. Data are
usually country- or region-specific, but trends in one region
may influence consumption in another region.
Food consumption data collected by governments,
academia, and industry monitor what has been eaten. If col-
lected in a consistent manner, they document changes in pat-
terns of food intake. These may also be useful in predicting
what will be needed in greater or lesser quantities in terms of
nutrition and the marketplace and may help distinguish
between what might be a fad and a trend.
Trend watchers and marketers conduct and scan material
from books, media, blogs, and other social media, especially in
areas that intersect with food such as cooking, health and
medicine, energy and the environment, employment and
income factors, and agricultural economics. Pronouncements
about nutrition by various authoritative bodies impact food
choices and messaging about food. Government regulation
also impacts what may be sold and what may be said about
it. For example, in Mexico, breakfast cereals are being rated
with a tax as a measure to address obesity. This causes cereal
manufacturers to reformulate to meet government guidelines
and to launch more high-fiber or reduced-sugar versions.
Understanding how these factors interact can help predict
consumer food trends.
Trends in Grain Usage and Its Impact on Consumers
Data on supply and disappearance of grains are followed in
many regions. For example, the Economic Research Service
(ERS) of the US Department of Agriculture (USDA) report
consumer demand for food by US households. This may
influence the types of crops that farmers grow; the prices
farmers receive; how crops are grown, handled, and
manufactured; and ultimately what grain products are
produced. ERS data on acres planted, yields, stores, and
other vital data for the major food grains (wheat and rice)
and the major feed grains (corn, barley, oats, and sorghum)
also give an idea of grain supply and stores to assess food
security for a population. The ERS working with the Census
Bureau and other government agencies. Publicly available
databases play a key role in assessment of nutrient intakes
and evaluation of the risks related to possible hazards in food
(Figure 1).
Trends in Worldwide Wheat Consumption
In the 1990s and the first decade of the 2000s, world wheat
consumption continued to expand in response to rising popu-
lation and incomes. Rising wealth fosters increased consump-
tion of meat, meaning that more wheat and grains are being
used as animal feed, thus boosting total usage. However, in
some countries, direct human consumption of wheat is
decreasing. For example, wheat consumption in the United
States has decreased since 1997. The pattern of wheat con-
sumption is shown in Figure 2. Per capita wheat use declined
from a high in 1879 when one in every four was on the farm
and many engaged in hard physical labor, making the per
capita consumption 102.27 kg. For nearly 100 years, until the
1970s, wheat use dropped to a low of 50 kg per capita as energy
required for both work and play decreased, wealth increased,
and diets became more diversified. By 1997, wheat use had
rebounded to 66.72 kg per capita. The overall growth in per
capita use that occurred between 1973 and 1997 reflected
changes that included (1) the boom in away-from-home eating
and fast foods with their buns or other grain-based food items,
(2) an increase in ready-to-eat options such as breakfast
cereals, (3) widespread dietary guidance that promoted diets
low in fat and high in carbohydrates and grains for controlling
coronary disease, and (4) a wider recognition of health benefits
stemming from eating bran, oat bran, and other high-fiber,
grain-based foods.
The 30-year pattern of increased wheat consumption ended
in 1997. There are many reasons for this trend: (1) the decrease
in energy expenditure, (2) concern about increases in rates of
overweight, obesity, and rising chronic disease, and (3) a num-
ber of missteps with the introduction of low-fat products, often
Production + beginning stocks + imports
minus
Exports + farm and industrial use + ending stocks
equals
US food consumption
(disappearance)
Figure 1 Estimation of food available for consumption (disappearance
data) in the United States. Data from USDA/Economic Research Service.
 FOOD GRAINS AND THE CONSUMER | Consumer Trends in Grain Consumption 31

U.S. per capita wheat flour use, 19642012

Pounds/person

150

140

130

120

110

100
1968 1972 1976 1980 1984 1988 1992 1996 2000 2004 2008 2012

Source: USDA, Economic Research Service calculations using data through the
second quarter of 2011 from U.S. Department of Commerce, Bureau of the
Census Flour Milling Products (MQ311A) and U.S. Department of Commerce,
Bureau of Economic Analysis Foreign Trade Statistic. Subsequent flour milling ERS
calculations are based on data from the North American Millers Association.

Figure 2 US per capita wheat flour use, 19642012.

formulated with grains, that failed to either address overweight
or add nutrient quality.
Thus, around the turn of the millennia, consumers
switched from low-fat diets to low-carbohydrate diets of vari-
ous types such as the Dr. Atkins Diet Revolution and the South
Beach Diet. Some of these ketogenic diets allowed only 200 cal
from all carbohydrates, so that there was a decrease in con-
sumption of all carbohydrates and grains. This switch reduced
per capita wheat and grain consumption. Consumer interest in
low-carbohydrate diets spiked after 2000, although this trend
continues because of medical literature showing that it is a
successful diet pattern for some dieters.
ERS estimated per capita wheat flour use at 60.22 kg in
2011. New allegations about wheat and wheat products from
blogs and books bashing gluten, wheat and wheat breeding,
and grains in general continue to keep wheat consumption at
lower levels than in 1997.
Whole Grains, Carbohydrate Quality and Glycemic
Response as Trend Drivers
The type of grain and wheat products is also changing. Since
2000, emerging science around the health benefits of whole
grains was translated into dietary guidance in many countries
with recommendations to substitute a portion of refined and
enriched grain staples with whole grain breads and cereals, for
example, to make half your grains whole. Across many
continents, this effort is reflected in groups such as Australia
and New Zealands Grains and Legume Nutrition Council or
the HEALTHGRAIN project in the EU. Nordic efforts to
increase rye and oats and other whole grains are part of a
healthy eating advice. In Italy and France, there also has been
a modest increase in whole grain products with successful
launches of whole grain pastas and other products. The
Whole Grains Council, which has international reach, has
documented a marked increase in the number of whole
grain foods and a shift to whole grain offerings in all
categories.
Data show that consumers in Western countries eat too
many refined grain servings and too few whole grain servings
such that their calories exceed their energy needs. Grain-based
desserts and snacks provide nearly as many calories as sugar-
sweetened beverages in the US diet of children and teens. Since
many of these grain foods offer little in the way of nutrition, it
is critical that health professionals and the food industry work
together to both improve the nutritional contribution of such
products and promote their consumption in the correct serving
sizes and as special treats.
Concern about the glycemic impact of foods including
wheat and grain products has caused some to use glycemic
index (GI) in as part of dietary advice, especially for diabetics.
Either some governments have allowed (Australia) GI labeling
or some merchants (the United Kingdom) have voluntarily
labeled foods including grain-based foods with GI. Some gov-
ernment bodies such as Health Canada have decided that the
GI is not appropriate for food labeling because of problems
with measurement and lack of consumer understanding. When
used, dietary guidance in these countries encourages con-
sumers to select products in the same category with GIs
32 FOOD GRAINS AND THE CONSUMER | Consumer Trends in Grain Consumption
deemed low to moderate. In some cases, foods carried a GI
label, and then, for a variety of reasons, the voluntary labeling
was withdrawn. Such labeling can impact the type of grain
foods selected and has become a driver in the selection of
many carbohydrate foods.
Gluten-Free and Wheat and Grain Avoidance Trends
Gluten-free and the avoidance of wheat, gluten, and grain have
become both a fad and a trend in some developed countries in
the last few years. The trend is also being exported and adopted
in some emerging economies.
Data published in 2008 changed the thinking in health
professions regarding the incidence of celiac disease. These
data showed that the incidence, instead of being 1 in 2500, is
around 1 in 133, and it is underdiagnosed. And data from the
United States, Finland, and even Asia Pacific countries where
the overall incidence is low show that celiac disease incidence
is increasing.
Nonceliac gluten sensitivity has been proposed as a gluten-
related disorder. The condition has no medically vetted diag-
nostic test and no agreement about its incidence, but is charac-
terized by a cluster of symptoms including gas and distention.
Beyond these conditions, the trend toward reduced wheat
and grain consumption is driven by books and blogs suggest-
ing that modern wheat and grains are addictive leading to
overconsumption of grains and are causing obesity and other
chronic diseases and health conditions. These same sources
demonize not only high sugar intake but also wheat and
grains, especially refined grains. Some make consumers believe
that foods labeled as gluten- or wheat-free are more nutritious
than their gluten-containing counterparts.
Summing of all those who need to avoid gluten those
with wheat allergy, celiac disease, and nonceliac gluten
sensitivity means that around 68% of the population
needs to avoid gluten. In 2014, as many as 30% of the North
American population said that they sometimes avoid wheat
and gluten. This is also a trend in Australia and New Zealand
with 10% of the population indicating that they are avoiding
gluten. In terms of markets, the United Kingdom has the
biggest gluten-free market in all Europe, but Germany has the
largest market on the continent. Italy and Scandinavia also
have a sizeable market. Yet, all of these countries have about
1% of consumers with celiac disease. Compared to other
European countries, sales of gluten-free foods are on the low
side in both France and Spain.
In terms of trends, the gluten avoidance fad will have an
impact for a while and will likely diminish. However, there will
be a baseline trend for those with celiac disease, allergies, and
other gluten issues that will remain and may increase. Concern
over carbohydrate quality will remain high, and changes in
types of carbohydrates and grain-based foods will serve as
another trend driver.
Rice
Rice supplies around 20% of the calories consumed worldwide
and is a staple for nearly half of the worlds 7 billion people.
More than 90% of the worlds rice is consumed in Asia, where
it is a staple for a majority of the population, including some of
the worlds poorest populations. Developing countries have
long depended and continue to rely on rices versatility, protein
availability, and high caloric value.
From the 1960s to the 1990s, Asias per capita rice con-
sumption went from 85 kg per year to nearly 103 kg. In that
same period, per capita consumption rose from 50 to 65 kg per
year. The rising per capita consumption plus the growing pop-
ulation more than doubled global rice consumption during
this period from 150 to 350 Mt.
Changes in the standard of living have altered rice
consumption patterns. Increased wealth in countries such as
India and China slowed the rate of growth in rice consumption
due to a switch to foods with higher value such as meat and
produce. In some countries such as Japan and Taiwan, there is
decreasing rice consumption due to Westernization of the
diets, a decline of family size, increasing participation rate of
women in labor market, and aging populations. However, rice
consumption continues to rise in Southeast Asia (the
Philippines and Indonesia), South Asia (India and Bangla-
desh), and many parts of Africa. After Asia, the second largest
rise in rice consumption occurs in sub-Saharan Africa. Overall,
the rising populations of emerging and developing economies
cause a rise in overall rice consumption. For those needing to
avoid gluten, rice has become an important substitute.
Recently, concern about arsenic in rice has been raised as an
issue and may impact consumption.
In summary, increasing wealth in some countries may shift
consumers away from rice and grains to other foods afforded
by higher-income consumers, but the increases in population
in many countries offset any decline in worldwide consump-
tion. Worldwide rice production was 600 Mt in 2000. It is
predicted to increase 1.5 times by 2030, to 904 Mt with gains
in production in nearly all areas.
Maize, Oats, Sorghum, and Barley
Maize (corn), sorghum, barley, and oats are used more as feed
grain than as human food. Maize contributes one-third of the
worlds cereal output. Demand for maize as a feed grain will
increase. Globally, maize production has increased by nearly
50% due to expansion in Asia and high-yielding varieties
including genetically modified corn (bT-corn). Differences in
consumer attitudes to the GM corn around the world have
dramatically affected its acceptance. The challenge this creates
for international trade of maize and future production must be
addressed and can sharply impact trends in its use as a feed or
food grain.
Only 15% of maize is processed for human food to become
a variety of milled grain products and starch, sweeteners, corn
oil, and alcoholic beverages. Direct use of maize as human
food is unlikely to change significantly, even in countries
where maize is an integral part of local diets.
Oats are an important foodstuff in northern climates
because they tolerate cooler summers and more rain than
many grains. Oat production has remained steady for the
past 10 years. The EU region is the largest producer of oats,
but Canada is the largest exporter. The United States is the
 FOOD GRAINS AND THE CONSUMER | Consumer Trends in Grain Consumption
worlds largest importer of oats, even though it is the worlds
fourth largest oat producer. Mexico is the worlds second larg-
est oat importer, followed by Japan.
Per capita consumption of oats is high in former colonies of
the United Kingdom with Canada and Australia topping the
consumption charts. Russia, the United States, Poland, the
United Kingdom, and Scandinavia are countries with a higher
consumption of oats than many countries. In the United
Kingdom, oats comprise 3% of the crops cultivated, but half
the oat crop is used as human food where about 50% of the
population eat porridge (oatmeal).
Among the grains, oats is viewed as a healthy grain. This
trend began with the oat bran craze of the late 1980s when
research documented the health benefits of oats. Its status as a
whole grain and high nutritional marks, with its micronutri-
ents, phytochemicals, and protein content ranking high among
the grains, continue to lure consumers.
The real driver for oats consumption is its viscous fiber
b-glucan. The data on its cholesterol-lowering ability are so
well documented that it has a health claim in a number of
countries. The first of many claims was in the United States in
1997 with other countries following. Interest in the health
value of oats is expected to continue as consumers continue
to want good-tasting, high-fiber whole grains. Emerging
research on benefits of oats for blood glucose control and
satiety keeps consumers interested, and the production of
gluten-free oats for celiacs is driving demand for oats.
The only concern is that oats is the only major world crop
where production has shown a steady decrease since 1970. The
decline amounts to a total of about 60%.
Barley
Only 2% of the barley crop is used directly for human food.
Malt production for beer takes over 40% of the crop. Barley,
like oats, contains the soluble fiber b-glucan, which has been
associated with maintaining healthy cholesterol levels and
healthy heart. In 2005, FDA amended its health claim for
soluble oat fiber and coronary heart disease to include barley.
Studies indicate that barley may also have potential in modi-
fying glycemic response in humans. Also, the vitamin E of the
outer layers of whole barley may be important. There is a trend
by consumers to romanticize ancient grains, and barley is
considered part of this ill-defined category. Use of barley as
sprouted grains is also showing some consumer interest.
Trends in Consumer Consumption
In many countries throughout the world, calorie intake has
increased since 1970. Despite charges to the contrary that
excesses are due to one food group such as grains, data from
ERS suggest that food availability and consumption for all
major food groups have increased. The ingestion of more
energy over the last three decades has not been translated
into more people meeting dietary recommendations for certain
food groups and nutrients.
The 2005 Dietary Guidelines for Americans recommend (for
those on a 2000-calorie-per-day diet) the consumption of 6
ounce equivalents (oz-eq) of grains per day with whole grains
33
accounting for at least half of this amount. Using ERSs loss-
adjusted food availability data from 2005, it is estimated that
more grain servings than recommended (8.1 oz-eq of grains
per person per day versus the 6 recommended) were con-
sumed. Instead of half of these as whole grains, nearly 90%
was as refined grains. Consumption data for children and teens
show that nearly as many calories emanate from consumption
of grain-based desserts and snacks as do from sugar-sweetened
beverages. Thus, Americans, on average, overconsume refined
grains with added sugar and fat and underconsume whole
grain staples. Only a small percentage of the population was
eating the recommended amount of whole grain. Efforts to
bring total amount of grain eaten in line with recommenda-
tions while substituting whole grains for refined grains must
continue in order to reverse current trends.
It is difficult to predict what economic and agronomic
conditions and policies, geopolitical climates, and technical
advances will affect food pricing and availability and costs
especially with population, fuel, and environmental pressures.
Trend drivers such as new health research, social media with
food fads and scares, changing social norms, adoption of foods
from other countries, and other issues related to consumers
will continue to affect consumption but are hard to predict.
Trend watchers and available data sources can track trends and
may be used to help answer critical questions about food
choices.
Summary
Consumption data for grains and grain-based foods relate to a
number of trends. Major factors that affect trends around the
world include demographics (age, household size, household
income, women in the labor force, amount of education,
geographic location, and ethnic background), diverse lifestyles,
and consumer attitudes (toward food habits and food prefer-
ences). Consumer trends that affect movement of grain-based
foods through the food system include food marketplace
trends, food processing/manufacturing trends, and agricultural
trends. Major trends in the food system include niche market-
ing, development of grocery stores to improve food shopping
convenience, increased demand for and development of con-
venience foods, and the growing demand for food away from
home.
Exercise for Revision
Compare the effects of agriculture policy in a country from
the developed world and developing world and see how it
affects grain consumption trends.
Follow two different grain food fads within a country and
see what caused the fad, how long they lasted, and at what
rate they rose and fell, and compare the drivers of the trend.
Compare a grain food fad in different countries and see
what is similar and different.
What are the trend drivers needed to change attitudes
regarding grains produced using biotechnology?
Examine the trend behind distrust of traditional breeding
among some sectors.
34 FOOD GRAINS AND THE CONSUMER | Consumer Trends in Grain Consumption
Exercises for Readers to Explore the Topic Further
Discuss impacts of inputs (water, fuel, fertilizer, etc.) on
trends in grain consumption in a region.
Study the interaction of food trends such as the locavore
movement, the Paleo diet, and other meat-based antigrain
diets and the organic movement on grain and food produc-
tion and the ability to feed a growing world population.
See also: Carbohydrates: Glycemic Index, Glycemic Load, and Their
Health Benefits; Health Effects of b-Glucans Found in Cereals; Food
Grains and the Consumer: Genetically Modified Grains and the
Consumer; Grains and Health: Misinformation and Misconceptions;
Grains and Health; Food Grains: Intolerance, Allergy and
Diseases: Celiac Disease; Grains Around the World: Grain
Production and Consumption: Overview; The Cereal Grains: Barley:
An Overview of a Versatile Cereal Grain with Many Food and Feed Uses;
Maize: Overview; Oats: Overview; Rice: Overview; Wheat: An Overview
of the Grain That Provides Our Daily Bread.
Further Reading
Abbassian A (2007) Maize international market profile. http://siteresources.worldbank.
org/INTAFRICA/Resources/257994-1215457178567/Maize_Profile.pdf (accessed
January 2015).
Economic Research Service (ERS), U.S. Department of Agriculture (USDA). Food
Availability (Per Capita) Data System. http://www.ers.usda.gov/Data/
FoodConsumption; http://ers.usda.gov/topics/food-choices-health/food-
consumption-demand/food-consumption.aspx.
Kubo M and Purevdorj M (2004) Future of rice production. Journal of Food Distribution
Research 35(1): 128142, http://ageconsearch.umn.edu/bitstream/27145/1/
35010128.pdf (accessed January 2015).
Lin BH and Yen ST (2007) The U.S. Grain Consumption Landscape: Who Eats Grain, in
What Form, Where, and How Much? ERR-50. U.S. Department of Agriculture,
Economic Research Service.
Mohanty S (2013) Trends in rice consumption. http://irri.org/rice-today/trends-in-
global-rice-consumption (accessed January 2015).
Rice Outlook/RCS-14l/December 12, 2014 Economic Research Service, USDA.
Wells HF and Buzby J (2008) Dietary Assessment of Major Trends in U.S. Food
Consumption, 19702005. Economic Information Bulletin No. 33. Economic
Research Service, U.S. Department of Agriculture.
World Agricultural Supply and Demand Estimates WASDE-537 January 12, 2015.
Relevant Websites
http://www.agr.gc.ca/eng/industry-markets-and-trade/statistics-and-market-
information/by-product-sector/crops/crops-market-information-canadian-industry/
market-outlook-report/oats-situation-and-outlook-august-2010/?
id1378843401619.
www.barleyfoods.org (accessed January 2015).
http://www.efsa.europa.eu/en/datex/datexfoodcdb.htm.
http://ers.usda.gov/data-products/food-availability-%28per-capita%29-data-system/.
aspx (accessed December 2014).
http://ers.usda.gov/data-products/food-availability-%28per-capita%29-data-system/
food-availability-documentation.aspx.
http://ers.usda.gov/topics/food-choices-health/food-consumption-demand.aspx.
http://www.ers.usda.gov/topics/crops/wheat.aspx.
http://www.ers.usda.gov/topics/crops/wheat/usda-wheat-baseline,-2014-23.aspx.
http://www.ers.usda.gov/topics/international-markets-trade/global-food-markets/
international-consumer-retail-trends.aspx.
http://irri.org/rice-today/trends-in-global-rice-consumption.
http://www.just-food.com/management-briefing/germany-canada-australia-among-
other-growth-markets_id123020.aspx.
http://www.nationmaster.com/country-info/stats/Agriculture/Grains/Oats/
Consumption-per-million (accessed January 2015).
 Cultural Differences in Processing and Consumption
AMR Hayes, St. Catherine University, Lafayette, MN, USA
JM Jones, St. Catherine University, Arden Hills, MN, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Consumption of cereal grain varies by cultural practice and
country, and each variety contributes unique nutrients that
can be affected by different processing techniques and prep-
aration methods.
The processing techniques used in different countries
impact starch digestion and energy yield of cereal grains;
most minimally processed grains are digested slowly, and
highly refined grains provide slightly more calories than
less refined types.
Cereal grains are incomplete sources of protein but can be
paired with other foods containing the amino acids they
lack to provide a complete source of protein, as is evident in
the traditional dishes of many countries.
Processing can either inhibit or promote the bioavailability
of minerals and toxic compounds, affecting nutrient needs
in certain regions.
Wheat is the most common cereal grain in the majority of
Western countries; refined or enriched flours are popular in
the United States, Europe, Latin America, South Asia, and
much of Australia.
Rice contributes approximately half of the calories for half
of the worlds population and is common in regions of
South and Southeastern Asia, Japan, Latin America, the
southern part of the United States, and the Philippines.
Discussions of cereal grains (teff (tef), oats, rice, corn/
maize, barley, rye, millets, sorghum, and triticale) and
pseudocereal grains (amaranth, buckwheat, and quinoa)
provide insight into their histories and current uses.
Increasing globalization and promotion of specific foods,
such as those that are gluten-free or whole grain (WG), are
altering the types and roles of cereal grains in the diet of
many regions.
Learning Objectives
To identify general variations in energy, protein, and nutri-
ent requirements among cultures.
To recognize and understand some of the changes in cereal
grains that occur with certain processing techniques or
cultural preparation methods.
To gain insight into the diverse histories of distinct cereal
grains, their nutritional qualities, and some of their current
uses.
To attain an understanding of cereal grains that promotes
the exploration for future applications addressing changes
in dietary habits.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00073-5
Cultural Differences in Processing and Consumption
Cereal grains are staple foods among many cultures. They can be
consumed separately or in combination with other foods. Each
variety of cereal grain contributes unique nutrients, mainly pro-
viding carbohydrates, protein, dietary fiber (DF), and vitamins
and minerals such as B vitamins, vitamin E, magnesium, and
iron. Additionally, different processing techniques can affect the
bioavailability of certain nutrients in cereal grains. The aims of
this article are to discuss the place of grains in the diet, the effects
of processing on grain nutriture, some specific types of grains,
and the overall impact of cultural practices of grain consumption
on diet.
Cultural Differences in Consumption
Energy requirements
Cereal grains can provide over 50% of both energy and protein
in parts of the developing world because they comprise the
fundamental components of most diets. Since nutrient require-
ments, food availability, and cultural practices vary by country,
the role of cereal varies. For example, abundant supplies of
food in developed countries result in cereals comprising a
smaller percentage of the diet and energy intake. The protein
contribution of cereal grains is less important for individuals
in the developed world than those in the developing world.
Exceptions may be for those who are vegetarian or flexitarian.
Cereal grains that are highly refined may contribute to more
available carbohydrate and slightly more calories than unre-
fined varieties. This is partly due to structures in the outer layers
of grain trapping some of the starch and making its form
resistant starch (RS) type 1. Germ and bran phytochemicals
may inhibit the activity of amylases and proteases, thus reduc-
ing the total number of calories available. High levels of tannin
in some grains such as tannin-containing sorghum varieties
can have a significant impact on calorie availability in coun-
tries where this grain is a staple. Refined grains may be bene-
ficial in developing countries, where calories are in high
demand. Alternatively, in developed countries where excessive
caloric consumption is widespread, the inclusion of WGs
with their slightly reduced carbohydrate contribution may
be advantageous.
Protein
Each cereal grain is characterized by distinct nutritional quali-
ties. When the cereal grain is a major food source in a culture or
region, its nutritional characteristics are of notable importance.
In developed countries, protein quality and quantity of grains
35
36 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption
are less important because other protein-rich foods are more
available and constitute a higher percentage of the diet.
Processing methods impact protein nutritional quality of
cereal grains. Milling, fermentation, and germination increase
the digestibility and bioavailability of protein and certain
essential amino acids. Germination also increases the synthesis
of the amino acids lysine and tryptophan. When cooking and
dehulling (debranning or decortication) accompany fermen-
tation of grains such as sorghum and millets, protein digest-
ibility is increased because antinutrients such as tannins are
reduced. In the case of tannins, this means that tannins bind-
ing of protein and inhibition of digestive enzymes (proteases)
are reduced.
Cereal grains are generally incomplete sources of protein, as
they are limited in the amino acid lysine. Corn (maize) is also
limited in tryptophan, while wheat, barley, and rye are low in
threonine. Through pairing cereal grains with other foods con-
taining the limiting amino acids, a complete protein can be
achieved. This protein complementation is exemplified
throughout the world by national dishes that pair cereal grains
with legumes. Cereal grains contribute methionine, which is
generally limited in legumes, while legumes contribute lysine
and other amino acids. Some regions use a grain-based dish
with a very small amount of animal protein such as cheese or
meat. The following are a few examples of popular dishes:
Ethiopia: Kik alicha wot (spiced cooked split peas) with
injera (teff flatbread)
India/Sri Lanka: Chapati (cereal unleavened flatbread) with
dhal (legumes, often lentils)
Middle East: Pita wheat bread with hummus (garbanzo
beantahini spread)
China: Congee (rice porridge) with toppings of tofu, fish,
and sometimes meat
Mexico: Corn tortillas, beans, and rice, sometimes with a
little meat or cheese
Southern Africa: Samp (porridge from coarse milled maize)
with whole common beans or cowpeas
The United States: Peanut butter on whole wheat bread
The United States: Macaroni and cheese
Carbohydrates, including DF
Starch
Energy from cereal grains is primarily from starch. Processing
impacts starch digestion and energy yield. When grain is milled
into flour, energy yield is increased. This is important in areas
of nutrient and energy deficiency. Germination and fermenta-
tion result in the partial degradation of starch, which promotes
starch digestibility. Grains that are cooked whole or are mini-
mally processed may be digested slowly, therefore slowing
glucose release into the bloodstream, referred to as slowly
available glucose (SAG). This may be advantageous for satiety
and control of blood glucose and lipid levels.
Processing can alter the ability of digestive enzymes to
access the starch. For example, the compact structure of an
al dente pasta impedes access of the digestive enzymes to the
starch, lowering the glycemic response (GR), but not as much
as is seen, for example, in a spatzle noodle from Germany.
With the porous structure of breads, even WG ones, amylases
have easy access to the starch and thus elicit a rapid GR.
Lowering the pH as occurs through a sourdough process
changes the activity of the amylase and lowers the GR. Cultural
practices of adding acid, as with oil and balsamic vinegar on
bread or vinegar in Japanese sushi, also lower the glycemic
impact. Using fats such as oils or butters on bread or using
legume and nut accompaniments also lowers the GR.
Resistant starch
Processing techniques can impact starch digestion. Processing
can affect the degree to which a starch is digested and absorbed
and can increase or decrease the content of RS. The following
processes increase RS: extrusion at high temperatures, as in the
making of cornflakes, followed by cooling to induce crystalli-
zation of the starch molecules. Thus, amylase enzymes do not
readily penetrate the starch, which means some starch is undi-
gested. Treatments of repeated heat and moisture, such as is
practiced in Africa where maize porridges are repeatedly heated
and cooled, also create RS (type 3). Cooked kernels and bul-
gurs, as is popular along the Mediterranean, trap starch in the
grain matrix and make it unavailable to digestive enzymes
resulting in RS (type 1). Microwave cooking shortens starch
chains in many foods and speeds digestion by having more
rapidly available glucose (RAG) and less RS. Germination and
fermentation not only may decrease RS but also may change
the levels of RAG and SAG
The following processes increase RS:
Steam cooking
Autoclaving
Parboiling
Baking
Extrusion cooking (some studies show it decreases it
slightly also)
Coarse milling
Cultural preferences for grain varieties or grain processing can
dramatically impact carbohydrate availability and the GR. For
example, the amount of RS and SAG in rice varies by processes
such as parboiling and variety. High-amylose varieties, such as
basmati preferred in India, have lower GR. Conversely, varie-
ties high in amylopectin, such as glutinous rice preferred in
parts of China and Japan, have high GR. Most forms of proces-
sing, notably parboiling, often reduce GR.
Dietary fiber
A growing body of evidence suggests that inadequate DF intake
is associated with decreased gut motility and increased risks of
heart disease, diabetes, and certain types of cancer. Diets in
many countries are characterized by inadequate intake of DF
and WG. Extraction rate, or the degree of milling, impacts the
nutritional quality of cereal grains. Higher rates retain more
bran and germ and therefore have a higher level of DF and
grain phytochemicals. Countries that regularly use higher
extraction rates, such as Norway, have increased DF and grain
phytochemical intakes.
DF intake is higher in countries that regularly include WG
in the diet. For example, regular consumption of WG products
made from whole wheat, rye, oats, and barley as occurs in
Scandinavia and parts of northern Europe contributes to
higher DF intakes than in many European nations. In parts of
 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption
Asia where white rice is the staple, the DF contribution of cereal
grains is frequently low. In a traditional rural Mexican diet with
its staple of WG corn tortillas and beans, DF intake is high,
whereas in the urban areas, DF may be lower due to a West-
ernized diet pattern.
The specific types of cereal grain typically consumed by
various cultures also affect nutriture, as each grain offers vari-
ous fiber types and phytochemicals. Rye, a common choice
among Germans and Scandinavians, is relatively high in arabi-
noxylans. Wheat, which is common in North America and
southern Europe, contains fewer arabinoxylans than rye, but
its cellulosic bran is associated with gut health and improved
laxation. b-Glucans, which are found in barley and oats, are
often consumed in northern Europe. In parts of the world
where high-tannin cereals such as sorghum are the staple,
nutrient availability, especially that of minerals, is compro-
mised. Additionally, some hull materials may be bowel irri-
tants or can damage the gut tissue (e.g., oat hulls with their
razor-sharp pieces).
Micronutrients
The bran and germ portions of cereal grain contain the major-
ity of the vitamins and minerals. Overmilling and high degrees
of extraction even as practiced in producing white bread flour
can remove the aleurone layers of grains, along with germ-rich
vitamins, minerals, and other grain phytochemicals. Thus,
limited extraction and the inclusion of WG foods as is part
of traditional cultural food patterns may help maximize
micronutrient amount contributed by grains. However, con-
sumption does not always equal absorption, as certain micro-
components may be bound or unavailable. While milling
reduces the overall number of micronutrients, it also makes
them more absorbable.
Heat treatment of some grains not only increases shelf life
but also is critical to preventing rancidity of all fats including
essential fatty acids, fat-soluble vitamins, and antioxidants
within the bran. Rice bran and oats are just examples of grain
or grain components that are typically subjected to heat treat-
ment in order to inhibit lipase activity of the grain and germ.
However, severe heat treatments may destroy vitamins, bind
nutrients, and potentially promote rancidity.
B vitamins in grains are particularly affected by processing.
Since thiamin is among the most heat-labile of the vitamins, it
is easily reduced through heating in alkaline pH. Thus, the
thiamin content of chemically leavened (baking soda) baked
products, such as Irish soda bread, can be decreased by over
70%. Conversely, processing via germination and fermenta-
tion can increase B vitamins. In the case of nixtamalization
(the process of liming corn), the tightly bound niacin is
released from niacytin to make it available to the body. Thus,
in countries where maize was traditionally limed, pellagra
(niacin deficiency) was not endemic.
Enrichment and fortification are used in many countries to
replace some of the vitamins and minerals lost during milling.
Such techniques are used to address nutrient deficiencies and
their corresponding health consequences. For example, pella-
gra was eliminated in the southeastern United States by niacin
enrichment of corn. In countries with mandatory folate forti-
fication of grain products, birth defects such as spina bifida are
reduced, in some cases, by over 50%. However, it is important
37
to note that enrichment and fortification efforts can be negated
by cultural practices. For example, washing rice prior to cook-
ing it or cooking it in large amounts of water, which is then
discarded, washes away much of the vitamin-rich talc added to
enrich the rice.
Fermentation and malting processes can alter nutrient qual-
ity. Fermentation typically results in increased levels of
vitamin B, but the extent of increase depends on the type
of grain and the particular process. Malting or germination of
grains and pulses has been found to increase vitamin C and
phosphorus availability. Such practices are especially impor-
tant in parts of the world such as India and Africa.
Mineral status is largely impacted by the grain choice and
processing of a certain culture. Ligands in grain, such as tan-
nins and phytates, bind iron, zinc, and other metals, rendering
them less bioavailable and leading to mineral deficiencies. How-
ever, absorption of some minerals can be improved through
processing techniques. Notably, milling with debranning
removes most of the tannin and some of the phytate, increasing
mineral bioavailability. The crushing and pulverization of the
grain may also increase bioavailability.
Traditional lactic acid (sourdough-type) fermentation can
facilitate the phytase reaction, releasing the tightly bound zinc
in many grains. The longer the fermentation process, the lon-
ger the phytase enzyme is allowed to degrade phytate, thus
improving mineral status. In the 1970s, hypogonadal dwarf-
ism was shown to be due to zinc deficiency, due to diets where
85% of the energy needs came from whole wheat pocket bread.
With efforts to increase the bioavailability of zinc in food and
improve overall diet, the incidence of this health condition has
been greatly reduced.
Toxic compounds
Aside from nutrient-binding phytates and tannins, cereals and
other plant foods can contain toxic or antinutritional sub-
stances. However, processing techniques such as milling can
remove a majority of these:
Protease and amylase inhibitors found in cereal bran are
lost or reduced during milling, germination, or fermenta-
tion processes. Grain soaking or heating as practiced in
some cultures may also reduce the activity of these
inhibitors.
Saponins that occur, for example, in some pseudocereals
not only can cause growth inhibition and are known for
their hemolytic activity but also may have anticancer
impacts on the colon. Again, the cultural practices of soak-
ing, fermentation, and germination may reduce these.
Mold toxins can affect any grain, especially those grown in
humid climates or harvested with high moisture levels.
Grain grading and inspection processes used in various
regions are in place to prevent various mycotoxins from
entering the food supply. Milling may remove mycotoxins
in the outer grain layers but not those contaminating the
endosperm. Industrial milling practices have steps that
remove unsound grains or incorrectly colored grains in
early points of the milling process as an attempt to remove
any infected grains. Fermentation and nixtamalization may
further reduce levels of aflatoxins, due to either an opening
of the lactone ring or a loss of the toxins into soaking liquid.
38 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption
Cultural Uses of Specific Grains and Their Impact
on Nutritive Qualities
Wheat and Its Parent/Derivative Grains and Products
Wheat
Wheat and its related grains such as durum spelt and other
wheat parents are used as flour, wheat berries, bulgur, cracked
wheat, and coarsely milled fractions such as farina or semolina.
All types of wheat can be used for pastas, but durum makes the
optimal product.
In most Western countries, more foods are made with
wheat than with any other cereal grains. While WGs are used,
refined or enriched varieties are popular in the United States,
Europe, Latin American, South Asia, and much of Australia and
Africa. Breads, pastries and small cakes, breakfast cereals, and
pastas are just some of the ways wheat is used. Noodles are
found around the world in various forms. In many countries,
they are made from durum wheat. Couscous, while often
mistaken for a grain, is a steamed semolina, which is a staple
of cooking in North African countries such as Morocco,
Algeria, Tunisia, Libya, and Egypt. To produce couscous,
yellow granules of semolina, which are made from durum
wheat, are parcooked and then dried. Its culinary uses as a
salad or pilaf base, an addition to soups, an accompaniment
for meats and vegetables, or a dessert make it like other small
grains and some pastinas. Both plain and flavored varieties of
couscous exist.
Bulgur
Whole wheat kernels that are broken to become cracked wheat.
If they are soaked, precooked, and then dried, they become
bulgur. Evidence indicates bulgur has been eaten by Arab,
Israeli, Egyptian, and Roman civilizations as early as
1000 BC, while the Chinese consumed bulgur as early as
2800 BC. Currently, bulgur can be sold as a pilaf or tabouli,
taboule, tabbouleh, or tabouleh mix.
Grano
Grano is a specialty durum that has been cultivated in Italy
since the fifth century BC. Its origins in Sicily and Puglia make
it a popular food choice on certain holidays, but it is a newly
emerging product in the United States. Firm and chewy in
consistency, grano is made from specially selected durum
wheat. Although it most resembles barley, its bran has been
removed, making the starch more accessible. Grano can be
cooked like arborio rice in dishes similar to risotto and either
used as a hot accompaniment with pasta or served cold in
salads.
Einkorn/farro/emmer
Einkorn and its cousin farro are hulled wheat. They were two of
the first types of wheat to be domesticated about 11 millennia
ago in Georgia, Armenia, and Turkey. Its name, however, is
from the German meaning first grain. Farro/emmer is known
by different names depending on the region, with the most
common ones being farro and emmer. Nutritionally, einkorn
is somewhat higher in protein than modern wheat or farro, but
all have lysine as their limiting amino acid. Einkorn contains
somewhat higher levels of phosphorus, potassium, iron, and
magnesium than wheat or farro. Like many ancient wheat
relatives, the lower yields and greater costs of einkorn and
farro make them less widely available. They can be used in
any way that wheat is used, and they are now being sought as a
specialty grain for variety.
Kamut
Kamut, a relative to durum (also known as khorasan), was
originally cultivated in the Fertile Crescent, and is still used
along the Mediterranean. Given its naturally higher sugar con-
tent, kamut has a sweet nutty flavor. It provides B vitamins,
selenium, and magnesium; is higher in zinc, magnesium, and
selenium than modern wheat; and provides a higher amount
of protein but a similar amino acid profile compared to hard
red wheat. Kamut can be used as whole kernels for pilafs or
ground into refined WG flour to be used in breads, pasta, and
other grain-based foods.
Spelt
Spelt is known as dinkel to Germans and farro (but different
from emmer, also called farro) to Italians. It dates back to 7000
BC. Like modern wheat, this ancient parent has winter and
spring varieties as well as hard and soft types. It is mainly used
in breads and pastas. Its protein content is only slightly higher
than modern wheat, but it is higher in selenium, phosphorus,
and magnesium than most cereal grains. Spelt WG flours and
doughs contain significantly higher amounts of RS and give
less available starch. However, baking or other processes affect
this, with breads showing the highest starch availability and
cookies the lowest. All of the grains related to wheat contain
gluten and must not be used in the preparation of products for
the gluten-free market.
Teff (Tef)
Teff, a tiny, slightly sweet-flavored grain, is so small its name
means lost grain. The bran and germ compose a larger per-
centage of the grain than other grains, and it has high levels of
RS. Teff provides fiber and B vitamins, has a higher overall
mineral content than most other cereal grains, and is known
for its levels of iron and calcium. However, it may not be as
high in iron as was once thought. Currently, it is also being
grown more widely in India, Australia, and Canada.
Traditionally, teff is ground into flour and fermented to
make spongy sourdough bread called injera (an Ethiopian
flatbread resembling a spongy large pancake). Teffs use in
some other countries has expanded into foods featuring
ancient grains or multigrain blends, such as tortillas, pizza,
crackers, pasta, pancakes, snacks, breads, cereals, and many
other products. Its application into gluten-free products is
expanding.
Oats
Oats are a popular grain in regions where they thrive, including
Russia, Canada, North America, Finland, Poland, and the
United Kingdom. In countries that were once part of the British
Commonwealth, porridge was standard breakfast fare with
steel-cut or pin-milled oats being preferred for their chewy
texture and nutty flavor. While this is still a breakfast option,
 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption
changing habits have caused a decrease in the intake of por-
ridge and an increase in oats in beverages, as additions to
yogurt and as the basis of ready-to-eat cereals and breakfast
and energy bars. For other parts in the British Isles and most of
Scandinavia, oats are found in many foods such as oatcakes,
biscuits, crackers, and even soup thickeners. In the United
States, most groats are steamed and flattened into rolled oats
(old-fashioned or regular oats, quick oats, and instant oats).
The higher the degree of flattening and steaming, the quicker
the cooking time and softness of the end oats. In Germany,
Switzerland, and other parts of Europe, muesli-type cereals,
based on uncooked, rolled oats with a combination of nuts,
fruits, and other grains, are popular.
Part of oats popularity is due to their nutrition and satiety
properties. They contain more than 20 unique polyphenols
called avenanthramides. They have high antioxidant, anti-
inflammatory, and anti-itching activities. They are high in
b-glucans. These fibers have been proven to lower serum cho-
lesterol and help maintain a healthy blood glucose level. A
number of countries have health claims that tout these benefits.
Rice
Rices roots date back to South Asia. It has been dispersed and
cultivated in a wide variety of habitats and consumed by
humans for centuries. It is grown on every continent except
Antarctica. Rice provides about half the calories for up to half
of the worlds population, especially in South and Southeast-
ern Asia, Japan, Latin America, the southern part of the United
States, and the Philippines. Rice has a role in religious and
ceremonial occasions and medicine in many cultures. It is so
important in certain regions that saying Hi. How are you? is
literally asking Hi. Have you eaten rice today?
Among various parts of the world, the type of rice desired
can be very different. Rice is classified by texture, color, and
size: long-, medium-, or short-grain varieties. Short-grain vari-
eties can be referred to as sticky because of their higher amy-
lopectin content (starch), while brown rice (WG rice) is known
for its higher levels of vitamins and minerals, and colored rices
are prized for their variety and antioxidant capacity, the latter
being due to their high anthocyanin content.
Among the diverse rice uses throughout the world are Chi-
nese stir-fries, Spanish paellas, Japanese sushi, Indian
biryani, Thai salads, Middle Eastern pilafs, Italian risottos, with the fourth highest production in the world. The amount
Mexican beans and rice, Senegalese yassa, Norwegian rice
pudding, American gumbos, and so many more.
Wild rice is not actually a kind of rice. Instead, it is a semi-
aquatic grass historically grown in lakes, tidal rivers, and bays.
Native Americans traditionally harvested the wild rice in north-
ern lakes and processed it by parching the green seeds in large
pots over open fires and then packing them away in birch-bark
baskets. Commercially, wild rice farmers cure the rice until the
chlorophyll dissipates, and then while the kernel is still in the
hull, it is dried. The inedible hull is removed mechanically,
leaving the kernel exposed. All these processes give wild rice its
signature nutty, smoky flavor. It is slightly higher in protein
than most other WGs and is a good source of fiber, folate,
magnesium, phosphorus, manganese, zinc, vitamin B6, and
niacin. Native Americans mixed it with berries and used it
39
with game. It is often mixed with other rice and can be used
any way that rice may be.
Corn/Maize
Corn, also known as maize, is a staple in parts of Central
America, Africa, India, and China. It can be used alone or in
combination with other foods and is the most widely grown
crop in the Americas. The domestication of corn occurred over
9000 years ago in Mexico. Corn was carried from the Americas
to Europe and Africa by early explorers and traders, so it
became important in the cuisines of many countries through-
out the world.
Maize contains a small amount of provitamin A (which can
contribute up to ten times the content of other grains) and is
high in antioxidants and carotenoids associated with eye
health (lutein and zeaxanthin). Its protein quality is poor and
niacin tightly bound, so it is associated with pellagra in mar-
ginal diets based on corn. Nixtamalization, a process used in
Latin America, releases the niacin and improves amino acid
availability in order to prevent pellagra. This process of soaking
the grain in lime (calcium oxide) gives it a flavor that charac-
terizes the Latin kitchen. Traditional dishes mixing corn with
beans and rice in many cultures provide amino acids to make a
complete protein. Flour from the nixtamalized corn is masa
harina, which is made into tortillas, enchiladas, and many
other dishes.
In the United States, corn was originally soaked in lye to
remove the pericarp. It was then dried to make hominy and
hominy grits. Hominy and corn meal are popular throughout
the southern and southwestern United States with foods such
as corn pone and hush puppies. Cheese and grits are popular in
the southeastern United States and spicier corn dishes and
hominy lamb stews (posole) are popular in the Southwest.
Corn-based breads and hot cakes are used throughout the
United States and can be made of corn alone or have wheat
added to them. The method and ingredients vary across the
country. In Italy and Spain, corn is used for dishes similar to
polenta. Ugali in Kenya and mealie porridge in South Africa are
just a few of the corn dishes that are also important in Africa.
Barley
Barleys use as human food is small despite its being the grain
of milling required to make the barley edible varies by whether
the grain is hulled or hulless (naked). Like chaff of the wheat,
the outermost layers are not digestible and must be removed by
processing. For hulless varieties, the outer hull is loosely
attached so very little polishing is required for its removal.
Whether hulled or hulless, most barley is polished (pearled)
to some degree, varying from regular to medium to fine to baby
pearled barley. On average, pearled barley is about one-third
lower in micronutrients than unpearled. Pot barley (also called
scotch barley) has the hull polished off, but the polishing
process is least invasive.
Barleys nutritional contribution is similar to wheat with a
few differences. It is highest in fiber of all the WGs, containing
on average 17 g of fiber/100 g, with a range from 12 to
40 g/100 g. The fiber in barley is b-glucans, and unlike in
40 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption
other grains, it is distributed throughout the grain. Barleys b-
glucans have a health claim in the United States and in other
countries.
Barley contains twice as many fatty acids as wheat, which
accounts for its slightly higher calorie content. Barleys lysine
content is higher than most other grains, giving it a better
amino acid profile. It is a gluten-containing grain.
Barleys prime use is for feed and beer, but it is also used for
making a tea in parts of Asia. Barley can be malted to make a
sweet syrup or addition to milk as in products such as milo or
malted milk powder. Additional applications include proces-
sing barley into flakes and other ready-to-eat cereals, porridge,
pancakes, and flatbreads; using it as pilafs and sides; and add-
ing it to hearty soups in areas of the British Isles to northern
and Central Europe.
Rye
Rye is grown in regions with colder climates, such as Russia, the
Baltics, parts of Germany, Poland, Scandinavia, Canada,
Argentina, China, and Turkey. Rye came to the Americas with
European settlers in the sixteenth and seventeenth centuries and
spread to South Africa and Australia during the mid-nineteenth
century. Like other grains, it is used to some extent in many
regions, but is most popular in northern and eastern Europe
where it has many uses and is popular in peasant-style breads.
Rye contains gluten, but much less than wheat, making many
rye breads heavy as in the classic German pumpernickel bread.
Rye mixed with wheat improves the texture and volume. Sour-
dough ryes are popular in a number of countries.
Rye berries may be soaked for several hours, added to a
small amount of sourdough rye culture, and baked; eaten hot
as a cereal; used in sides such as pilaf; added to soups; or used
in a grain salad. Rye can be flaked or added in other forms to
ready-to-eat cereals, and thin crisp rye breads are very popular.
Millets
Millet actually refers to several different small-seeded cereal
grain species, the major ones being pearl millet, foxtail millet,
proso millet, finger millet, and teff (although the latter is
usually considered separately). Millets are thought to have
been the staple grain in Asia before rice, so their cultivation is
over 10 000 years old. They are an important food source in
many parts of the world and are used widely in developing
counties. They are generally drought- and temperature-
tolerant, making them well suited for cultivation in semiarid
tropics in Asia and Africa, especially in India, China, Nigeria,
and Niger. Millet seeds are small, round, and relatively soft, of
varying colors and are generally rich sources of antioxidants.
Finger millet is notably rich in calcium.
Millets can be eaten as a breakfast porridge in regions such
as Africa and China. They can be added to soups and stews,
popped as corn, or made into sides such as pilafs and couscous.
Millets are used in breads, breakfast cereals, cookies, and crack-
ers. Millets are gluten-free grains so they used for foods in this
growing market. In India, ragi (finger millet) is used to make
roti (staple flatbread). In Africa, millets are also used for brew-
ing beer and in China, for wine.
Sorghum
Sorghum originates from Africa around 8000 BC. It thrives in
adverse conditions, making it versatile and popular. Sorghum
is known by many names milo or guinea corn (West Africa),
mabela (South Africa), dura (Sudan), mtama (eastern Africa),
jowar (India), or kaoliang (China). Since sorghums hull (actu-
ally pericarp) is edible, it can be eaten with all its outer layers
and thus retains the majority of its nutrients. WG sorghum
contains essential B vitamins, minerals, fiber, antioxidants,
other phytonutrients, and policosanols. The latter are thought
to impact cardiac health.
In Africa and parts of Asia, sorghum is used primarily as a
human food. The light color and neutral flavor of white tan-
plant (the so-called food-grade) sorghum varieties easily
enable their substitution for wheat flour, wheat-based, and
gluten-free and multigrain baked goods including muffins,
breads, pizzas, pastas, casseroles, cookies, cakes, and pies. Sor-
ghums gluten-free, non-GMO WG status has increased its use
in the West. It works well in gluten-free baking mixes, cereals,
and breads and provides a WG option. Region-specific appli-
cations include the following:
North Africa: couscous and flatbreads
Bangladesh: boiled like rice to produce kichura
Honduras: used to make tortillas
Sub-Saharan Africa: porridges and traditional beers
India: flatbreads
China: distilled spirits and vinegar
Triticale
A hybrid of wheat and rye, triticale is barely over one century
old. It offers the advantages of wheat for product development
and the hardiness of rye to thrive in colder climates. Since it
requires few pesticides, reduces soil erosion, captures soil nitro-
gen, and capitalizes on the nutritional advantages of its wheat
and rye parents, it may be important in the future. It can be
used in most applications that its parent grains can be used.
Pseudocereal Grains
These are actually the seeds of dicotyledonous plants, but are
starch-rich, like cereal grains.
Amaranth
Amaranth not only was a staple grain of the Mayans and Aztecs
in Mexico but also has historical importance in Africa, India,
and Nepal. It has been grown in China, Russia, Thailand,
Nigeria, and parts of South America. Its name originates from
the Greek word amarantos, meaning one that does not wither
or the never-fading.
This tiny grain contains enough lysine to blend it with other
grains to improve protein quality. It contributes iron, phos-
phorus, magnesium, potassium, and folic acid, but it is noted
for its calcium contribution. Thus, this gluten-free grain is a
welcome addition to traditional and gluten-free formulations
for cereals, breads, muffins, crackers, and pancakes to add
 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption
additional texture and flavor. Some amaranth applications are
listed as follows:
South American countries: sold on the streets (like
popcorn)
India, Mexico, Nepal, and Peru: traditional ingredient for
breakfast porridge
Mexico: dulce de alegria, a sweet cookie-confection dessert
Buckwheat
The origins of buckwheat can be traced to about 8000 BC in
Tibet and northern China. Despite wheat in its name, it is in
the same family as rhubarb and sorrel but its seeds are used and
cooked as cereals are. The triangular-shaped, dark-hulled groat
and flour made from the entire groat are considered WG.
Buckwheat is used to make everything from pancakes to noo-
dles. Buckwheat noodles are important in Tibet, northern
China, Japan, and Korea and northern Italy as pizzoccheri. The
fact that these noodles contain no gluten makes their prepara-
tion and manufacture different from noodles made with wheat.
Roasted buckwheat groats, kasha, are popular in eastern
Europe, particularly Russia and Poland, where it is used as a
pilaf, mixed with pasta, added to filling for knishes and blin-
tzes, and used in soups and other dishes. Sprouted groats can
be eaten raw or cooked as porridge. Delicate buckwheat pan-
cakes (blini), sometimes raised with yeast, are eaten in several
countries, while heartier buckwheat hotcakes are eaten in
North America. In Ukraine and other regions, yeast rolls are
made from buckwheat. In Nepal, a doughy mixture of buck-
wheat with grains such as maize and millet called dhedo is
popular. A buckwheat, wheat, and maize mixture makes a type
of polenta eaten in northern Italy.
Buckwheat is a source of higher-quality protein than cereal
grains and is high in magnesium and potassium. It is gluten-
free, so it is also used in these applications.
Quinoa
Quinoa was a sacred staple food of the Incas. It is related to
beets and spinach. Over 120 varieties exist, but the white, red,
and black types are commercialized. Most of the quinoa con-
sumed within the United States is imported from South
America.
Quinoa is gluten-free, provides high-quality protein, and
contains potassium, folic acid, and vitamin E. Its bitter
saponin-containing outer coating offers natural protection
against pests, aids in drought resistance, and is an antioxidant.
It has a distinctive taste with mild notes of corn and bean
flavor. Frequently added to pilafs, stews, salads, or bread,
quinoa is found in gluten-free products and gourmet food
applications.
Conclusion
The variety of ways in which cereal grains are processed and
consumed around the world is both interesting and important
in terms of providing calories and contributing to nutrients to
the diet. The extent of grain refinement and cultural practices
41
determines the presence and bioavailability of micronutrients
and macronutrients. In most developed countries where
overweight is a problem, consumption of WGs may be
advantageous, whereas refined grains remain popular and
may benefit areas where calories are in short supply. With
increasing globalization, grains and foods specific to one
region are spreading across the globe and enriching the diets
of many both nutritionally and culturally. The resurgence of
plant-based and gluten-free diets has made traditional and
alternative cereal grains important dietary contributors.
Exercises for Revision
In one continent, take a sampling of counties from various
sectors and outline the main grains used and characteristic
dishes that show protein complementarity.
Compare a population where one staple grain is used with
another where a different grain is used and show the advan-
tages and disadvantages for each area.
If northern climates get hotter and dryer, discuss how grain
staples would change and suggest some cultural adapta-
tions of customary dishes in the region to be prepared
with these alternate grains.
Show how milling practices in two regions affect nutrition
outcomes in these regions.
What are the cultural issues that cause resistance to chang-
ing preferred grain and processing techniques?
How can various grains impact DF intake? If the diet is
based on a grain with a low-fiber content, what are some
dietary strategies to address this?
Some components in certain grains are antinutritional or
other concerning components. How can processing and
cultural practices minimize the impact of these?
Exercises for Readers to Explore the Topic Further
With an increasingly globalized world, how are the tradi-
tional cultural uses of cereal grain changing?
How can cultural preparation methods and practices be
altered to improve diet?
How can processing techniques and cultural preparation
methods be applied to provide optimal nutrient availability
in cereal grains?
What implications do the expanding gluten-free and WG
markets have on cultural practices?
Discuss how alternate grains can be utilized in diets that are
gluten-free or wheat-free. Cross-reference to gluten-free,
sorghum, millets, teff, and quinoa.
Show how WG inclusion in a diet where the cultural prac-
tice to choose refined grain can improve nutrient intake.
Cross-reference to WGs; see specific grains.
See also: Carbohydrates: Cereals: Chemistry and Physicochemistry
of Non-starchy Polysaccharides; Food Grains and the Consumer:
Consumer Trends in Grain Consumption; Processing of Grains:
Starch: Uses of Native Starch.
42 FOOD GRAINS AND THE CONSUMER | Cultural Differences in Processing and Consumption

Further Reading Shewry PR (2007) Improving the protein content and composition of cereal grain.
Journal of Cereal Science 46(3): 239250.
Abdel-Aal E and Rabalski I (2008) Effect of baking on nutritional properties of starch in
organic spelt whole grain products. Food Chemistry 111(1): 150156.
Jones JM (2012) Glycemic index: The state of the science, Part 1 The measure and its
variability. Nutrition Today 47(5): 207213.
Molina J, Sikora M, Garud N, et al. (2011) Molecular evidence for a single evolutionary Relevant Websites
origin of domesticated rice. Proceedings of the National Academy of Sciences
108(20): 8351. http://www.grdc.com.au/
Mosley BS, Cleves MA, Siega-Riz AM, et al. (2009) Neural tube defects and maternal http://www.homebaking.org/hbamembers/
folate intake among pregnancies conceived after folic acid fortification in the United http://hsc.csu.edu.au/food_technology/issues/
States. American Journal of Epidemiology 169(1): 917. http://www.ifst.org/learninghome/helpforteachers/lessonplantopics/cereals/
Schroeder N, Gallaher DD, Arndt EA, and Marquart L (2009) Influence of whole grain http://www.kswheat.com/
barley, whole grain wheat, and refined rice-based foods on short-term satiety http://wbc.agr.mt.gov/
and energy intake. Appetite 53(3): 363369. http://dx.doi.org/10.1016/j. http://www.wheatfoods.org/
appet.2009.07.019. http://wholegrainscouncil.org/whole-grains-101/
 Fortification of Grain-Based Foods
CM Rosell, Institute of Agrochemistry and Food Technology (IATA-CSIC), Paterna, Valencia, Spain
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Fortification is a way of supplementing the diet through the
intake of some nutrients to combat deficiencies.
Foods selected as vehicles for fortification must meet cri-
teria of availability and price.
Cereals are the best carriers for food fortification, mainly in
the case of vitamins and minerals.
Chemical compounds must be carefully selected for ensur-
ing micronutrient bioavailability after processing.
Fortification is legislated in different countries.
Learning Objective
Understand the meaning of fortification and the role of
cereals and cereal-based foods in fortification.
Introduction
Fortification can be defined as the addition of one or more
vitamins and/or minerals to a food, regardless of their usual
content in the food. Additions are carried out based on gener-
ally accepted scientific knowledge of the role of vitamins and
minerals in attaining good health. Foods are fortified for the
following reasons:
to prevent or correct a demonstrated deficiency of one or
more vitamins and/or minerals in the population or spe-
cific population groups; and
to improve the nutritional status of the population and
dietary intake of vitamins or minerals due to changes in
dietary habits.
Enrichment is a term usually interchanged with fortification.
The term enriched flour was coined during the World War II
when the American government decided to improve the
soldiers diet by including different nutrients in bread. Initially,
the enrichment consisted of the addition of the three major
B-vitamins (thiamine, riboflavin, and niacin) and iron, despite
the scarce scientific information concerning the real nutritional
improvement obtained through that proposal. Later, numer-
ous scientific studies have been conducted for verifying the
utility of those measures.
Ideally, enrichment should be equivalent to restoration.
This means that the addition of vitamins and minerals, present
in the edible portions of the food, will restore the levels lost
during manufacturing, storage, and handling.
The other relevant term in this subject is nutritional
equivalence, which means being of similar nutritive value in
terms of quantity and bioavailability of vitamins and minerals.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00074-7
Since fortification is a broader term, it will be used here while
referring to the addition of nutrients.
Criteria for Food Fortification
Minerals and vitamins may be added to foods only for the
purpose of (1) restoration of nutrients lost during processing,
(2) nutritional equivalence of substitute foods, (3)
fortification, and (4) ensuring the appropriate nutrient com-
position for a special purpose food.
Some considerations should be taken into account prior to
addition of any nutrient to foods:
The nutrient added should be present at a level that will not
result in either an excessive or an insignificant intake, con-
sidering the amount provided from other dietary sources.
The maximum safe amount to be added should be set
considering the levels established by generally accepted
scientific data and bearing in mind the diverse degree of
sensitivity shown by different consumer groups.
Nutrients should not be added to fresh products including
meat, fish, fruits, and vegetables, neither to beverages con-
taining more than 1.2% by volume of alcohol.
The nutrient added should be biologically available from
the food and should not interfere with the metabolism of
any other nutrient.
The nutrient added should be sufficiently stable during
packaging, storage, distribution, and end use and should
not affect both the sensory characteristics and the shelf life
of the food.
Technology and processing facilities should be available to
permit the addition of the nutrient and also methods of
measuring and controlling the levels of nutrient added.
Fortification should not be used as a tool for providing
nutrients in substitution of the natural nutrient sources, as
this would lead to a change in the dietary patterns and thus
jeopardize good dietary practices.
It is clear that the policy for food fortification has a completely
different objective when considering developed and develop-
ing countries. In developed countries, the social changes
(industrialization, high per capita income, consumption of
fast food outside the home) over the years have led to increas-
ing consumption of more highly processed foods. This has
been used to justify the necessity of food fortification for
ensuring the adequate intake of nutrients, mainly vitamins
and minerals. However, there is no general consensus about
the extent to which food fortification should be allowed. There
are differing opinions that manufacturers could use fortifica-
tion as a promotional tool, resulting in an excessive intake of
certain nutrients that would represent a risk for consumer
43
44 FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods
health. An uncontrolled food fortification program could
result in excessive intake of certain nutrients, thus creating
nutrient imbalances, and in consequence representing a risk
to the health of consumers. It must be had in mind that while
deficiencies provoke diseases, the intake of too much of a
vitamin or mineral also may cause health problems. In fact,
too much vitamin A can cause birth defects, too much folate
can hide symptoms of a vitamin B12 deficiency which can lead
to damage of the nervous system, and too much calcium can
induce kidney problems.
In developing countries, food fortification is one approach
toward solving nutrient deficiencies, and so far the implemen-
tation of these programs has been very successful in correcting
nutritional deficiencies of large segments of the population
within a very short period. In these countries, care should be
taken that the food selected as a medium for the nutrient
addition is stable and routinely consumed by the population
at risk, and that the amount of nutrient added is sufficient to
correct the possible deficiency.
However, in any case fortification programs should include
adequate Quality Assurance and Quality Control (QA/QC) at
production level, regulatory monitoring of the nutrient content
of fortified foods, as well as assessment methods to document
the nutritional and health effectiveness when implementing
fortification strategies.
Cereal-Based Products as Vehicles of Minerals
and Vitamins
One of the most critical decisions concerning food fortification
is the selection of the appropriate food to act as a carrier of the
nutrient. The basic criteria for identifying the food products for
fortification include selecting a food that is regularly consumed
by the target population group, affordability, and availability
all through the year.
Cereals and cereal-based products constitute a major com-
ponent of diet around the world. Cereals have been used as the
basis of human diet from the ancient times and their implica-
tion in the human diet is still very significant.
Globally, cereals account for 48% of total energy intake and
they are a cheap source of proteins compared to animal pro-
teins. Although their relative contribution to human diet
greatly depends on the production and the cultural differences,
the popularity of cereals, mainly rice and wheat, in the diet
worldwide make them the most appropriate vehicles for forti-
fication strategies. Additionally, it must be remarked that
cereals are not directly consumed as grains; processing is gen-
erally required to transform the grain prior to its consumption,
with the subsequent reduction in nutrients and micronutrients
that are mainly concentrated in the outer layers and germ. In
the conversion of cereal grains to white flour, 20 or more
ingredients (such as thiamin, riboflavin, niacin, pyridoxine,
folate, panthothenate, biotin, vitamin E, calcium, copper,
iron, potassium, magnesium, manganese, phosphorus, zinc,
chromium, fluorine, molybdenum, and selenium) are lost to
the extent of 5090% during the milling process. Cereal forti-
fication methods have been developed to restore the nutrients
that have been removed during milling and to improve the
nutrient intake level of specific population.
Table 1 Vitamins and minerals that may be added to foods
Vitamins Minerals
Vitamin A Calcium
Vitamin D Magnesium
Vitamin E Iron
Vitamin K Copper
Vitamin B1 Iodine
Vitamin B2 Zinc
Niacin Manganese
Pantothenic acid Sodium
Vitamin B6 Potassium
Folic acid Selenium
Vitamin B12 Chromium
Biotin Molybdenum
Vitamin C Fluoride
Chloride
Phosphorus
Once the food or carrier to be fortified is selected, the
nutrient deficiencies affecting the population must be deter-
mined. There are only certain vitamins and minerals that may
be added to foods (Table 1). Nevertheless, the main micronu-
trients recognized as being of concern to the health of the
population are iron, folic acid, B-vitamins, vitamin A, and
zinc, although in developed countries calcium also is
considered.
In general, the population of all the rice-consuming coun-
tries have vitamin A deficiency, which is associated with cor-
neal lesions that can lead to partial or total blindness, and with
reduced resistance to infectious diseases. This would, in conse-
quence, lead to an increase in morbidity and mortality.
Another deficiency associated with people of rice-consuming
countries is anemia due to iron deficiency. This has been linked
to reduced resistance to infection and also affects the cognitive
development and physiological functions in children and, in
severe cases, causes maternal deaths. Iodine constitutes the
third major deficiency in rice-consuming countries. This min-
eral is necessary for proper fetal development and also for
normal physical and mental activities in adults.
In developed countries, iron supplementation has been a
measure adopted to reduce the prevalence of iron-deficiency
anemia. B-vitamins (thiamine, riboflavin, and niacin) and
calcium are also supplemented in developed countries to
ensure the adequate intake of those nutrients. Lately, some
studies confirming the role of folates in congenital malforma-
tions (neuronal tube defect) and the development of chronic
diseases like Alzheimers in elderly people have recommended
the supplementation of folic acid in cereal-based foods. How-
ever, the effectiveness of this fortification program in the
decrease of homocysteine concentration and its relationship
with a reduction in the incidence of cardiovascular disease is
still under research.
Selenium is an essential micronutrient for humans and may
reduce the risk of degenerative diseases, including cancer, but
its deficiency has been detected in at least a billion people
worldwide, although wheat is a major source of dietary sele-
nium in humans.
 FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods
Methods for Cereal Fortification
Cereal fortification methods are mainly addressed to restore the
nutrients that have been removed during milling and to improve
the nutrient intake level of specific population. In addition, some
efforts have focused on enhancing the nutritional quality of the
crops because half of the world population suffers from malnu-
trition with respect to iron, zinc, and selenium.
Biofortification
Biofortification, the increase of micronutrients in the edible
parts of the plants, can be achieved by either mineral fertiliza-
tion or plant breeding. Biofortification by mineral fertilization
is a common practice in some countries that applied selenium-
containing fertilizers as a short-term solution for improving
selenium content of wheat.
Biofortification through plant breeding consists of the
development of micronutrient-enhanced crop varieties
through conventional breeding. Nevertheless, the implemen-
tation of this type of biofortification requires the identification
of genetic resources to determine the interaction of genotype
and environment, to define the desired level of micronutrient
increase, and finally to assess the cost-effectiveness. So far, it
seems a sustainable and cost-effective approach for reaching
biofortification in zinc and iron.
Rice Fortification
Rice requires special methods for fortification as it is com-
monly consumed as a whole grain and is extensively washed
prior to cooking or strained after cooking. There are different
methods for rice fortification; the earliest one was the produc-
tion of parboiled rice. In the parboiling process, rice is steamed
under pressure to gelatinize the starch within the kernel; this
process allows the transference of nutrients from the bran layer
to the starchy endosperm.
Other methods currently used are the powder and grain
enrichment methods. A powder preblended with mixtures of
vitamins and minerals is added to the rice. For white parboiled
rice, the blend is added immediately after milling because the
heat and moisture content of the grain surface at that moment
facilitates the adherence of the powder mixture. However, the
Table 2 Average levels of nutrients to consider adding to fortified wheat flour based on extraction, fortificant compound, and estimated per capita
flour availability
Level of nutrient to be added (ppm) by estimated average per capita wheat flour availability
(g day
1)
Nutrient Flour extraction rate Compound <75
Iron Low NaFeEDTA 40
Ferrous sulfate 60
Ferrous fumarate 60
Electrolytic iron Not recommended
High NaFeEDTA 40
Source: WHO, FAO, UNICEF, GAIN, MI, and FFI (2009) Recommendations on wheat and maize flour fortification. Meeting Report: Interim Consensus Statement. Geneva: World Health
Organization. http://www.who.int/nutrition/publications/micronutrients/wheat_maize_fort.pdf (accessed 10.12.13).
45
nutrient supplementation in this method can be washed off if
the grains are rinsed before cooking.
The grain enrichment method involves the addition of a pow-
dered mixture of vitamins and minerals and a subsequent coating
of the grain with a water-insoluble substance. Usually, grains
enriched by this method contain a high nutrient concentration,
and they are mixed in a proportion of 0.5% with normal milled
rice to finally obtain an enriched product that meets the required
standard levels. Different variations of these methods have been
developed through the years. For instance, the powdered mixture
can be sprayed onto the rice and then coated with ethanol,
isopropanol, palmitic acid, or cellulose derivatives. This method
has been effective for enriching rice with niacin, thiamin, pyri-
doxine, vitamin A, vitamin E, folic acid, iron, and zinc by adding
alternative layers of nutrients and coatings.
Synthetic rice obtained by extrusion of rice flour to a rice
kernel shape in the presence of vitamins and minerals also
covers the fortification objective. The fortified simulated grains
are then mixed with normal milled rice, although the con-
sumer often notices and removes those grains before cooking.
This is obviated by dyeing them with a distinct color, so that
the consumer clearly knows that it should be present.
Flour and Bakery Products
Fortification of flour is usually carried out in mills, where the
nutrient mixture is blended with the flour. In this practice, the
definition of the nutrients and the levels to be added must be
based on the nutritional needs and deficiencies of the popula-
tion, the common consumption of the fortifiable flour, the
sensory and physical aspects, and economic cost. The addition
of B-vitamins, iron, and calcium is a common practice in some
developed countries. However, the nutrient added must have
good stability during storage, and in defining the levels to be
added must be considered the extraction rate during milling. In
fact, some recommendations have been established for wheat
flour fortification (Table 2).
Fortification can even take place during the breadmaking
process. Different yeast strains have been suggested for obtain-
ing enriched baked goods. A selenium-enriched yeast has been
used for increasing the content of selenium in bread, obtaining
wheat bread rolls that can provide 25% of the recommended
daily allowance in the form of selenomethionine, which is the
best form of selenium for human consumption.
75149 150300 >300
40 20 15
60 30 20
60 30 20
Not recommended 60 40
40 20 15
46 FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods

Pasta Fortification Table 3 Compounds that may be added to cereals as a source of

vitamins
In pasta-consuming countries, pasta and noodles are used for
fortification. The production of fortified pasta or noodles Vitamins Chemical compound
requires only the addition of the nutrient at the mixing stage
prior to the dough extrusion process, or instead previously Vitamin A Retinol
Retinyl acetate
fortified flour can be used for this purpose. Some vitamin
Retinyl palmitate
losses are produced during the drying process, although the
Beta-carotene
extent of that loss depends on the temperature reached during Vitamin B1 Thiamin hydrochloride
drying. Thiamin monohydrate
Vitamin B2 Riboflavin
Riboflavin 50 phosphate
Breakfast Cereal Fortification Niacin (B3) Nicotinic acid
The fortification of ready-to-eat breakfast cereals is a well- Nicotinamide
Pantothenic acid (B5) D-Pantothenate, calcium
established practice. In fact, this plays an important role in
D-Pantothenate, sodium
ensuring the adequate intake of nutrients by the US
Vitamin B6 Pyridoxine hydrochloride
population. Pyridoxine 50 phosphate
Vitamins, like niacin and riboflavin, are added to the basic Pyridoxine dipalmitate
ingredients prior to processing when the breakfast cereal is heat Vitamin B12 Cyanocobalamin
stable or sprayed onto the processed breakfast cereal in the case Hydroxocobalamin
of addition of heat-labile vitamins (vitamin A, C, and thiamine), Vitamin C Ascorbic acid
ensuring the uniformity of the coverage. Hydrophobic vitamins Sodium ascorbate
(e.g., vitamins A and D) need special consideration as emulsions Calcium ascorbate
need to be sprayed on the cereals. In these products, the packag- Ascorbyl 6-palmitate
Vitamin E Tocopherol
ing material is very important, as the moisture content should be
Tocopheryl acetate
below 5% to avoid the texture modification of the product.
Tocopheryl acid succinate
Folic acid Pteroylmonoglutamic acid

Compounds Used as a Source of Vitamins and Minerals

Three different requirements should be considered when
selecting the source of nutrients: the chemical compound (1)
should be stable during packaging, storage, distribution, and
end use; (2) should not modify the sensory characteristics
(appearance, texture, taste, and aroma); and (3) should ensure
the bioavailability of the nutrient. Different chemical com-
pounds are used as a source of vitamins (Table 3) and minerals
(Table 4) to be added to cereals.
Nutrient stability is very important in the case of vitamin
fortification. Vitamins are sensitive to heat, oxidizing and
reducing agents, light, and other physical and chemical stress.
For instance, vitamins are stable when added to flour but the
combination of high humidity and temperature adversely
affects the stability of the vitamin A. This vitamin is labile to
high temperature and unstable in the presence of oxygen and
light, due to its susceptibility to oxidation. Therefore, a protec-
tion and appropriate conditions of storage should be used in
order to reduce the vitamin A loss. Encapsulation in a more
hydrophilic coat is a common practice in order to obtain a
more water-dispersible product.
The supplementation of B-vitamins to flour is affected by
low stability of these compounds during storage. Hence, higher
effectiveness is obtained when B-vitamins are added at the
bakery rather than at the mill.
The bioavailability of added nutrients, particularly iron,
varies widely depending on the iron compound used for forti-
fication and the physical and chemical properties of the food
to be fortified. The selection of the iron form is based on
organoleptic considerations, bioavailability, cost, and safety.
Iron powders are extensively used in flour fortification, but
they are poorly absorbed and their absorption is highly depen-
dent on the meal served (phytates, calcium, and polyphenols
decrease the bioavailability of iron). Ferrous sulfate is preferred
for enriching wheat flour because of its high bioavailability but
it often leads to unpleasant colors and flavors due to reactions
with other components of the food matrix. Some of these
interactions with other food components can be avoided by
coating the iron form with hydrogenated oils or ethyl cellulose.
Iron fortification of flour is generally regulated in most coun-
tries by specifying a minimum iron level in the flour resulting
from the sum of the added and native iron. Nevertheless,
Hurrell et al. reviewed the effectiveness of the iron fortification
programs in 78 countries and concluded that they are likely to
be ineffective. Legislation needs to be updated in many coun-
tries so that flour is fortified with adequate levels of the recom-
mended iron compounds to meet population requirements.
Legislation Concerning Cereal Fortification
Although cereal fortification can be an effective means of
improving micronutrient status, there are some barriers to the
general implementation. These include concerns related to a
proliferation of the fortified foods if the fortification is volun-
tarily practiced by manufacturers, which would lead to a simul-
taneous replacement of the nonfortified foods in the diet. The
success of fortification programs always depends on good con-
trol, so they should be set up, regulated, and enforced by the
national governments.
 FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods 47

Table 4 Compounds that may be added to cereals as a source of There are currently 14 countries with legislation or regula-
minerals tions that mandate the fortification of wheat flour, whereas in
countries such as Finland, New Zealand, and Norway law pro-
Minerals Chemical compound
hibits the addition of nutrients to wheat flour. In the United
Calcium Calcium carbonate States, the Food and Drug Administration approves the supple-
Calcium chloride mentation of four B-vitamins (thiamine, niacin, riboflavin, and
Calcium salts of citric acid folic acid) and iron to the wheat flour, for obtaining enriched
Calcium gluconate flour, although this practice is regulated by Federal law. The US
Calcium lactate government adopted a mandatory regulation concerning folic
Calcium hydroxide acid fortification. In the United Kingdom, it is mandatory for
Copper Cupric carbonate the restoration of all the nutrients removed with the bran in all
Cupric citrate
types of flours, with the exception of whole meal. From 1997,
Cupric gluconate
the Organizacion Panamericana de la Salud (OPS) is support-
Cupric sulfate
Iodine Sodium and potassium iodide ing the supplementation of iron to wheat flour due to its low
Sodium and potassium iodate cost and extensive consumption. Some countries from South
Iron Ferrous carbonate and Central America have implemented this measure for
Ferrous citrate reducing the incidence of iron-deficient anemia. The levels of
Ferric ammonium citrate nutrients to be added are also regulated by national
Ferrous gluconate governments.
Ferrous fumarate At the international level, GAINs (Global Alliance for
Ferrous lactate Improved Nutrition) National Fortification Program, which
Ferric diphosphate
began in 2003, has expanded to support 19 countries with
Elemental iron (carbonyl electrolytic hydrogen
high levels of vitamin and mineral deficiencies. A broad coali-
reduced)
Hydroxocobalamin tion of governments, businesses, international organizations,
Magnesium Magnesium acetate and civil society partners, known as National Fortification
Magnesium carbonate Alliances, support GAINs projects. Those projects aim to for-
Magnesium chloride tify foods and condiments including wheat and maize flour,
Magnesium salts of citric acid sugar, vegetable oil, milk, soy sauce, and fish sauce. Vitamins
Magnesium oxide and minerals used to fortify foods include vitamin A, vitamin
Magnesium sulfate D, iron, zinc, folic acid (B9), thiamin (B1), riboflavin (B2),
Manganese Manganese carbonate niacin (B3), pyridoxine (B6), and cobalamin (B12).
Manganese sulfate
Manganese chloride
Manganese citrate
Phosphorus Calcium salts of orthophosphoric acid
Zinc Zinc acetate
Zinc chloride Future Prospects
Zinc oxide
Zinc sulfate Currently, in the developed countries the term functional
Zinc carbonate foods or nutraceuticals seems to be old-fashioned, and
the cereal-based products surely may be one of the best
carriers of different compounds related to physiological
functions. Therefore, the near future in nutrition could be
The addition of nutrients is generally practiced by manu- the use of cereal-based products as carriers of o-fatty acids
facturers either voluntarily or because it is compulsory under and probiotic compounds such as fructo-oligosaccharides,
national rules. In general, vitamins and minerals may be added inulin, etc.
in conformity with the legislation of the country in which the
product is being sold, which must regulate the list of foods to
which vitamins and minerals may be added, the vitamins and
minerals permitted, and the levels at which they may be added. Exercises for Revision
The Economic and Social Department of FAO has elabo-
rated a review about the legislation status concerning rice What requirements should the nutrients to be used in
(Table 5) and wheat flour (Table 6) fortification. fortification meet?
Rice fortification is voluntary in the United States; however, When and where would food fortification be recommended?
most of the rice sold is fortified, and lately folic acid is also Which cereals would be the selection for food fortification?
being added to the enriched rice. In Canada, fortification of When using wheat white flour as a fortification carrier,
rice is also voluntary, and when labeled as enriched, it contains which chemical compounds would be advisable to be
vitamin B1, vitamin B3, and iron. Moreover, in Canada, the added? Cross reference to wheat milling chapters.
addition of vitamins B5 and B6 and folic acid is optional. Which cereal-based foods are usually fortified?
Fortification of rice in Philippines has a long tradition, and Considering iron deficiencies and consumption intake, jus-
there is a mandatory regulation for the addition of vitamin B1. tify where would be recommended iron food fortification.
48 FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods

Table 5 Summary of voluntary and mandatory fortification of rice

Country Law status Legislation Known nutrient deficiencies

Angola No Vitamin A, I, Fe
Australia Prohibited
Canada Voluntary B1, B3
Costa Rica Voluntary No I, Fe
Cuba Voluntary No Fe, vitamins B1, C, and A
El Salvador No Vitamin A, I, Fe
Ethiopia No Vitamin A, I, Fe
Finland Prohibited Restoration allowed
Gambia No I, Fe, Ca, folic acid, vitamin B2
Haiti No
Hungary Voluntary 1 serving must contain 1/3 of RDA
Mauritania No I, Fe, folic acid
Morocco Voluntary No Vitamin A, I, Fe
New Zealand Prohibited In process of reviewing the legislation
Norway Prohibited
Peru Voluntary No
Philippines Voluntary Legislation for B1, B3, Fe Vitamins A, B1, and B2, I, Fe
Sweden May be allowed with special permission
Tanzania No Fe, I, folic acid, vitamin A
Turkey Voluntary In accordance with Codex Fe, I, Ca, vitamins B2, B6, C, and D
United Kingdom Voluntary Must be safe
Uruguay No Fe, I, vitamin D
United States Voluntary B1, B2, B3, folic acid
Vietnam Voluntary No Vitamin A, I, Fe

Table 6 Summary of voluntary and mandatory fortification of wheat flour

Country Law status B-vitamins Vitamin E Folic acid Iron Zinc Magnesium Calcium

Australia Voluntary B1, B2, B3, B6

Canada Mandatory B1, B2, B3
Voluntary B5, B6
Chile Mandatory B1, B2, B3
Costa Rica Mandatory B1, B2, B3
Dominican Republic Mandatory B1, B2, B3
Ecuador Mandatory B1, B2, B3
El Salvador Mandatory B1, B2, B3
Finland Prohibited
Guatemala Mandatory B1, B2, B3
Honduras Mandatory B1, B2, B3
Hungary Voluntary
Malta Voluntary B1, B2, B3
New Zealand Prohibited
Nigeria Mandatory B1, B2, B3
Norway Prohibited
Panama Mandatory B1, B2, B3
Philippines Voluntary
Saudi Arabia Mandatory B1, B2, B3
Sweden Voluntary B1, B2, B3, B6
Switzerland Voluntary B1, B2, B3
United Kingdom Mandatory B1, B3
Uruguay B1, B3
United States Mandatory B1, B2, B3
Venezuela Mandatory B1, B2, B3
 FOOD GRAINS AND THE CONSUMER | Fortification of Grain-Based Foods
Exercises for Readers to Explore the Topic Further
The cereal composition changes after milling. What would
be the minerals and vitamins that must be restored after
milling? Cross reference to cereal processing chapters.
Select some cereal-based goods and determine which chem-
ical compound would be advisable for their fortification in
minerals.
Select several countries and identify their regulations
regarding fortification of wheat flour.
See also: Barley, Rice and Maize Processing: Barley: Milling
and Processing; Rice Processing: Beyond the Farm Gate; Food
Grains and the Consumer: Cultural Differences in Processing and
Consumption; Labelling of Grain-Based Foods; Food Grains:
Intolerance, Allergy and Diseases: Nutrition: Beriberi, A
Deficiency Related to Grains; Grain Composition and Analysis:
The Composition of Food Grains and Grain-Based Products; Grains
Around the World: Grain Production and Consumption: Overview;
Processing of Grains: Cereals: Breakfast Cereals; Wheat
Processing: Wheat: Dry Milling; Wheat-Based Foods: Breads.
49
Further Reading
Allen L, Benoist B, Dary O, and Hurrell R (2006) Guidelines on Food Fortification with
Micronutrients. Geneva: World Health Organization and Food and Agriculture
Organization of the United Nations. http://www.who.int/nutrition/publications/
guide_food_fortification_micronutrients.pdf (accessed 10.12.13).
Baurenfiend JC and DeRitter E (1991) Foods considered for nutrient addition: Cereal
grain products. In: Baurenfiend JC and LaChance PA (eds.) Nutrient Additions to
Food, pp. 143209. Connecticut: Food and Nutrition Press.
Hurrell R, Ranum P, de Pee S, et al. (2010) Revised recommendations for iron
fortification of wheat flour and an evaluation of the expected impact of current
national wheat flour fortification programs. Food and Nutrition Bulletin 31
(1 supplement): S7S21. http://www.ncbi.nlm.nih.gov/pubmed/20629349.
Ranum P, Lynch S, Bothwell T, et al. (2001) Guidelines for Iron Fortification of Cereal
Food Staples. Washington, DC: SUSTAIN. Published online at www.sustaintech.
org/publications/pubm7.pdf.
Rosell CM (2008) Vitamin and mineral fortification of bread. In: Hamaker B (ed.)
Technology of Functional Cereal Products, pp. 336361. Cambridge: Woodhead
Publishing Ltd.
Rosell CM (2012) The nutritional enhancement of wheat flour and related products.
In: Cauvain S (ed.) 2nd edn., Bread-Making: Improving Quality, 2nd edn.,
pp. 687710. Cambridge: Woodhead Publishing Ltd.
WHO, FAO, UNICEF, GAIN, MI, and FFI (2009) Recommendations on wheat and maize
flour fortification. Meeting Report: Interim Consensus Statement. Geneva: World
Health Organization. http://www.who.int/nutrition/publications/micronutrients/
wheat_maize_fort.pdf (accessed 10.12.13).
 Genetically Modified Grains and the Consumer
A Mathiowetz, University of Minnesota, St. Paul, Rochester, MN, USA
JM Jones, St. Catherine University, Arden Hills, MN, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The purpose of genetically modified (GM) grains is to
improve yield, increase resistance to plant pests and diseases,
help plants adapt to stressful environmental conditions,
confer resistance to certain herbicides, reduce spoilage mech-
anisms, and improve the nutrient profile of a crop.
GM grains may help address nutritional deficiencies in the
developing world.
The scientific community agrees that GM foods present no
greater safety risk than conventionally grown crops.
Despite prevailing scientific evidence, consumers remain
wary.
Awareness of GM foods is increasing among consumers, as
is skepticism too.
European, Mexican, and some African consumers are the
most outspoken against GM foods, while consumers in the
United States are the most open to the idea.
Consumer perception is dependent on understanding of
safety, perceived ratio of risks to benefits, religious/moral
values, intended use of GM foods, scientific literacy, trust in
regulatory organizations, price, and social economic status.
Yield gains through GM technology are critical for the
developing world to meet growing population, but mone-
tary benefits often occur in developed countries.
Maize, soy, and rice are commercially grown and sold
worldwide, but GM wheat has not been commercialized.
Labeling is mandatory on all GM foods and foods contain-
ing GM ingredients in 21 EU countries, Australia, Brazil,
Russia, China, and Japan. Individual states in the United
States are considering GM labeling.
Learning Objectives
To understand the purpose of GM grains and their applica-
tion to the greater majority of the world.
To recognize common factors that influence consumer per-
ception of GM grains.
To compare and contrast consumer perception of GM
foods/grains across varying regions of the world and to
observe differences in the perceptions of the scientific and
consumer communities.
To understand the movement for labeling of GM foods:
Where is labeling already mandated, who are its propo-
nents/opponents, and what is the fundamental argument?
Introduction
Genetically modified (GM) foods and grains create varied reac-
tions around the world. Some see the use of this technology as
50 Encyclopedia of Food Grains, Second Edition
a way to feed the rapidly rising population against a back-
ground of significant pressure on land, water, and other valu-
able resources. At the other end of the spectrum are those who
think that the use of this technology is playing God and have
dubbed GM foods frankenfoods.
GM grains refer to those that contain a gene introduced by
recombinant DNA methods to confer a desired trait. The abil-
ity to pick a specific gene for a desired trait is an outgrowth of
standard breeding practices that have been used for centuries
where plants or animals with desired traits were bred together
in hopes of perpetuating the trait. For many grains, this would
be a 10-year process. The use of genetic modification, also
called biotechnology, has been used since the 1970s. It can
shorten the time required to achieve the same outcome as
standard breeding process and can be much more precise
than traditional hybridization. Traits have been introduced to
grain foods to improve yield, to increase resistance to plant
pests and diseases, to help adapt to stressful environmental
conditions such as drought or extremes of temperature, to
confer resistance to certain herbicides, to reduce spoilage, and
to improve the nutrient profile of a crop.
Since the introduction of commercially available biotech
crops in 1996, the global use of GM crops have sustained a
growth rate of 6% or 10 million hectares per year. Thus,
between 1996 and 2012, there has been nearly a 100-fold
increase in acreage. Farmers in North America grow the most
GM crops with Brazil in second place. GM soybeans, maize,
cotton, and rapeseed were planted on 170.3 million hectares
worldwide in 2012, according to figures from the International
Service for the Acquisition of Agri-biotech Applications
(ISAAA). In developing countries, biotech grains accounted
for 52% of all biotech crops planted in 2012 worldwide. This
is up from 50% 1 year earlier. In contrast, Europe grew < 0.1%
of the worlds biotech grains.
Scientific bodies around the world continue to evaluate
evolving data and continue to aver that there are no health
risks in consuming foods including grains that are produced
by biotechnology. The US National Academy of Sciences
concluded no adverse health effects attributed to genetic
engineering have been documented in the human
population. Similarly, the International Council for Science,
which includes 111 national academies of science and 29
scientific unions, concluded that there is no evidence of any
ill effects from the consumption of foods containing GM
ingredients. The World Health Organization also issued a
response stating, No effects on human health have been
shown as a result of the consumption of such foods by the
general population in the countries where they have been
approved. In 2010, a European Commission review of 50
studies on the safety of biotech crops found no scientific
evidence associating GMOs (Genetically Modified Organisms)
with higher risks for the environment or for food and feed
http://dx.doi.org/10.1016/B978-0-12-394437-5.00075-9
 FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer
safety than conventional plants and organisms. The American
Medical Associations Council on Science and Public Health
stated that Bioengineered foods have been consumed for close
to 20 years, and during that time, no overt consequences on
human health have been reported and/or substantiated in the
peer-reviewed literature.
Despite acceptance by farmers and diverse expert scientific
panels around the world, consumer attitudes to GM foods and
grains vary markedly by geographic region. Various degrees of
activism, prevailing public opinion, and the extent of eco-
nomic development strongly influence attitudes toward GM
foods. However, two overarching factors seem to influence
public opinion of GM grains worldwide: (1) the riskbenefit
perception related to the personal and family situations and
needs of the country and (2) ambivalence or fear about the
safety of GM foods, often resulting from either a lack of infor-
mation or misinformation generated by some inaccurate
media. Often, media reports fuel fears of some untoward con-
sequence that may not be accurately predicted. In terms of risk
and benefit, some feel that the consumer bears a dispropor-
tionate share of the risk while large companies reap the mon-
etary benefit. In terms of adopting biotech crops, the farmers
are as divided as public opinion. Subsistence farmers worry
about planting biotech crops because they fear they may be
unable to feed their families or earn a living. In contrast, other
farmers are lured by the promise of fewer inputs of water,
fertilizer, and pesticides and thus lower costs and higher yields.
The view is strongly influenced by region.
Perceptions of GM Grains by Region
GM Foods in the United States and Canada
Farmers in North America have dramatically increased their
planting of GM grains between 1996 and 2013. This has been
driven by farmers expectations of higher crop yields and/or
lower production costs, a reduction in time devoted to crop
care and management, and other benefits including farm sus-
tainability. Reduced input, lower use of pesticides and herbi-
cides, less acreage and tilling required, and fewer crop diseases
are among some advantages cited. As a result, currently 93% of
the soybeans and 80% of the corn, according to the United
States Department of Agriculture (USDA), grown in the United
States are GM. (Despite allegations to the contrary, no GM
wheat is grown commercially in any part of the world.)
Consumer acceptance of GM food in the United States has
increased slowly but steadily and somewhat parallels the
increase in number of hectares of biotech crops. It also reflects
a less disparaging reaction to the technology by the press and
the public. This is reflected in the International Food Informa-
tion Councils 2012 survey on Consumer Perceptions of Food
Technology, which shows that 74% of consumers have, at
minimum, heard a little about biotechnology and as many
as 38% of US consumers are somewhat or very favorable
toward plant biotechnology. 44% asserted they were
somewhat or very favorable regarding the use of biotechnol-
ogy in grains specifically. Overall, consumers were likely to be
more accepting biotechnology if the benefits of the technology
were clearly communicated and especially if consumers were
51
convinced that the technology could increase yields for devel-
oping countries.
Variability in consumer perception is influenced by demo-
graphics of the consumer. Older consumers are more likely to
accept the use of biotechnology that affords health benefits,
not as likely to accept it for benefits of increased freshness or
flavor. Those with more education are more likely to try GM
crops. Women are more resistant to the use of biotechnology
than men.
Canadians are also divided regarding the concept of GM
foods. Less than a third of Canadian consumers say they
oppose the use of GM in food. This is a change in attitude
with more respondents in 2011 stating that they strongly
support or somewhat support the use of GM in food produc-
tion than in 2001.
Regulation in various countries often reflects public opin-
ion. Health Canada regulates GM foods and requires that
developers show that the seeds and plants have no untoward
effect on the environment or human health. Receiving
approval from Health Canada can take 10 years and be very
costly. However, once approved, Canada does not require
labeling on all GM foods, unless there are any health or safety
concerns such as a food containing an allergen.
The United States currently only has voluntary labeling
guidelines for foods produced by biotechnology. However,
the Food and Drug Administration (FDA) outlines certain
circumstances where labeling is required, including foods
that have significantly altered nutritional makeup, allergens,
or toxic levels of an ingredient as a result of GM. Regulatory
policies on GM foods are enforced by the FDA, Environmental
Protection Agency (EPA), and USDA.
The federal law regarding GM voluntary labeling does not
reflect the results of several polls indicating that over 90% of
US consumers want GM foods to be labeled. This is reflected in
bills on GM labeling being introduced at the state level.
Approximately half of US states have pending legislations
requiring labeling of GM foods to some extent. Maine and
Connecticut have passed laws mandating labeling of all GM
foods, and many states are likely to follow suit. Those in favor
of labeling argue that they have a right to know what is in their
food, some for ethical or religious reasons and some for health
or environmental reasons. Much of the argument for labeling
is simply transparency. Americans claim a right to have imme-
diate access to information on their foods content, so they can
make a personal decision.
Organic foods cannot contain GM ingredients, so some
consumers used the organic label as a proxy for a label saying
GMO-free. Many manufacturers and sellers of organic foods
oppose the use of GM products. This meets the wishes of many
organic consumers, but lack of GM labeling supports their own
economic interests.
The calls for legislation on labeling of GM foods have
dismayed those in agribusiness as well as some scientists and
consumers. Considering that 6070% of processed foods in
the United States contain at least one GM ingredient, experts
worry that the reaction of consumers to such labels will cause
them to reject use of GM products and seeds, turning back
gains made in yields along with profits. Food economists
suggest that not only it is these large corporations that will be
burdened with the negative consequences of labeling but also
52 FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer
consumers would likely see increased prices and changed avail-
ability. Groups who do not favor labeling often reference
scientific literature, arguing that there is no significant differ-
ence in the safety of GM foods and non-GM foods. They also
note that manufacturers are required to alert consumers of any
material difference in GM foods that could affect safety such as
the presence of allergens or potential toxins or decreased nutri-
tional quality. Those who oppose GM labeling state that con-
sumers wishing to avoid GM ingredients have the option to
purchase foods labeled organic, which precludes their use.
GM Foods in Europe
Strong resistance to GM foods pervades Europe. This occurs
despite numerous decisions regarding the safety of GM foods
by bodies such as the European Commission (EC). However,
regulators dealing with GM foods in Europe invoke the pre-
cautionary principle, which states that if there are any unre-
solved concerns, they will not accept the food process or
additive in the food supply. This principle aims at minimizing
any perceived or theoretical threats. It fails to consider any
potential benefits where the science is indeterminate. It also
is problematic as safety cannot be proven, only lack of safety.
Consumer resistance to GM foods and belief in the precau-
tionary principle have led at least 21 European Union (EU)
countries to require labeling of GM foods or foods with GM
components.
The overall sentiment in Europe is that the benefits must
outweigh the risks. Furthermore, consumers and experts hold
wholly different viewpoints toward GM foods, specifically
regarding safety, ethical tradeoffs, benefits, and international
trade. What might be seen as a benefit to experts is often viewed
as a risk to consumers. When GM foods were first introduced
into the European market, consumers did not perceive that
there was a benefit of these products in that they were not
cheaper, did not seem tastier, and did not offer other added
value. The higher yields per hectare did not reduce prices for
consumers due to a general increase in prices for fuel and other
inputs that affect food prices.
Food scares in Europe in the 1990s eroded confidence in
food regulatory bodies and the food supply. This influenced
discussions regarding the acceptability of GM foods. Con-
sumers continue to question aspects of consumer health, envi-
ronmental risks that may occur over time, and economic effects
on small European farmers. There are also fears of allergenicity
and the development of microbial resistance to GM traits just
as has occurred with pesticides and herbicides.
Farmers are wary of GM food as most of the GM food is
imported, so there is an economic concern as well. Countries
who produce GM products have contested the use of the
precautionary principle as a trade barrier. Yet organizations
such as the World Trade Organization are reluctant to impinge
on the authority of various member states.
Despite over 25 years of use and no environmental or other
problems, European consumers claim they have insufficient
and inconclusive information. They state that their confidence
could be improved if more definitive scientific information be
made more readily available to the public.
Acceptance of GM foods varies among the countries with
some of the strongest resistance found in the United Kingdom.
2013 surveys carried out by the UK government showed that
only 21% of the UK public support GM foods and less than
half of UK farmers believe GM technology is a good innova-
tion. Over 40% of people stated that they were completely
against the government promoting GM technology.
European acceptance of GM food, while limited, may be
increasing. A poll of European consumers conducted between
2002 and 2005 showed an increase of 6% in positive attitude
toward GMOs. 80% of EU consumers did not intentionally
discriminate against GM foods when shopping. Thus, Euro-
pean consumers stance on GM food may not be affecting
purchasing behavior. Recent research indicates that acceptance
of GM food is much improved if the consumer benefits such as
produced with fewer sprays and higher levels of a needed
component such as omega-3 fats are clearly communicated
to the consumer.
Asia-Pacific Nations-Developed Countries
In Australia, consumer acceptance of GM foods has fallen since
2007, following a steep rise in consumer acceptance between
2005 and 2007. However, the acceptance level among the
general public is still at 67%. Approval ratings of biotechnol-
ogy in food and drink have consistently remained lower than
approval ratings of biotechnology in medicines, vaccines, and
stem cells, as well as for the purpose of creating pest-resistant
plants. Awareness of biotechnology in food is at a commend-
able 82%, and a solid two-thirds of the Australian public thinks
the use of biotechnology in food crops could be useful. Less
than half of the population finds the application risky, but
GM-containing foods must be labeled.
A study of 2000 New Zealand consumers sought to deter-
mine whether consumers would purchase GM foods. Slightly
more than 30% of the over 700 who returned the survey said
that they would not purchase any of the GM foods listed on the
survey, which included bread from GM wheat and maize
altered to resist insects. However, not all information from
New Zealand shows attitudes toward it to be negative, espe-
cially when the science is applied sensibly and consumers see it
involves no great risks. Further biotechnology would be accept-
able in New Zealand if (1) the consumer is given a choice, (2)
the benefits hold greater weight than the risks, (3) the benefits
are shared by the consumer as well as agribusiness, (4) the
practice is environmentally friendly, and (5) the technology is
safe, practical, and cost-effective. Concern and reasonable
doubt certainly exist in New Zealand regarding GM foods,
but not to an extent that completely stifles acceptance.
Japan and Brazil, Russia, and China mandate labeling of all
GMO-containing foods. While Japanese resistance to GM is not
as strong as that in Europe, Japanese consumers continue to
remain unwilling to accept GM products. This was recently
highlighted after finding of GMO wheat in Oregon. This
revived the issue and Japans Ministry of Agriculture, Forestry
and Fisheries opted for temporary import suspension and
called for urgent greater GM inspection on imports to Japan.
Asia-Pacific Nations-Developing Countries
In Malaysia, almost 90% of consumers surveyed were aware of
GM foods. However, the fraction of Malaysian consumers who
 FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer
accept GM foods may not be as high. The study revealed large
concerns over food safety and perceived riskbenefit ratio.
Other factors affecting consumer opinion were media influ-
ence and lack of information.
Filipinos, in general, trust the international agencies
responsible for conducting studies and disseminating informa-
tion on topics such as GMOs. This may be true due to the work
of the International Rice Research Institute (IRRI) and the work
of the International Maize and Wheat Improvement Center
(CIMMYT). Citizens are less confident in the government
than university scientists. Overall, Filipinos believe the nutri-
tive qualities of GM grains could prove beneficial, but they
remain guarded about safety. A study conducted in the Philip-
pines analyzed the attitudes of 423 Filipinos, who worked in a
variety of careers. The majority were middle-aged, educated
males with an allegiance to Roman Catholicism. Attitudes
were affected by education level and religious affiliation;
those with higher education held more of a positive view on
GM foods, while the strongly religious had moral concerns
about GM technology. Many Filipinos across various disci-
plines claimed insufficient information on GMOs. Those who
received information from the scientific community were more
likely to have positive opinions of the technology than those
who received information from popular media. Even without
easy access to literature on biotechnology, Filipinos are less
likely to have extreme attitudes toward GM food. However, an
anti-GMO group of 400 interfered in 2013 with a GM trial on
Golden Rice, a variety of rice intended to decrease the preva-
lence of vitamin A deficiency.
India
In India, most consumers are aware of GM crops, some to a
greater extent than others. While a whopping 96% of Indian
consumers deem biotechnology to be beneficial and worth
further exploration, < 33% currently support the implementa-
tion of GM crops and hardly any want GMOs in their own food
supply. Although awareness is high and an overwhelming
majority views them as a promising prospect for the future,
consumers do not necessarily trust the use of GMOs in their
food currently. They are more open to their use in medicine.
The Supreme Court in India has just called for an indefinite
moratorium on field trials for new GM food crops. Just next
door, Bangladesh is allowing the planting of GM staples.
China
Chinese consumers remain divided over the use of GM foods
but are more accepting than consumers in Japan. A study
published in 2010 by Zhang et al. recognized Chinese con-
sumers as belonging to one of four categories: neutral (risks
posed by GMOs are equal to benefits), relaxed (benefits of
GMOs outweigh the risks), skeptical (risks outweigh the ben-
efits), and uninterested (no perceived risks or benefits). Social
status and income level strongly influenced the particular cat-
egory. Although variability in degree of acceptance exists across
China, results of the Zhang study suggest that more people
accept biotechnology than object to it. When asked to rate
nutrition-improved GM rice on a scale from completely
acceptable to strongly opposed, 25% of 1000 surveyed said GMO use, South Africa is the most progressive, and GM crops
53
it was completely acceptable, 42% deemed it relatively
acceptable, 21% pleaded neutral, 5% were relatively opposed,
and only 1% were strongly opposed. The acceptance rate of
folate-enhanced rice was found to be > 60% in a study of
nearly 1000 consumers. Again, results were most strongly
affected by perceived riskbenefit ratio.
Africa
Attitudes regarding GMOs in many countries in Africa are
influenced by those in the rest of the world. Some developed
nations and world food organizations encourage the use of GM
crops for the purpose of alleviating hunger and tackling food
insecurity. Not all Africans are convinced that GMOs would be
a helpful strategy in addressing food insecurity. In fact, many
are distrustful despite a backdrop of decreased agricultural
productivity throughout most of the continent. The Center
for Strategic and International Studies (CSIS) noted that
despite 70% of Africans working in agriculture, Africa imports
a quarter of its food.
Across Africa as a whole, the mood toward GM foods can be
summarized as ambivalent, with some nations more resistant
and others more welcoming. Tanzania, Uganda, Malawi, Mali,
Zimbabwe, Nigeria, and Ghana have at least begun to consider
the possibility of growing GM plants, and many are conducting
research and field studies.
Awareness and acceptance vary dramatically across Africa.
For example, the awareness level for GMOs in Ghana has been
calculated at around 60%, meaning more than half of the
population of Ghana has heard to some extent of GM foods.
However, increased awareness does not assuage fear; rather, it
may lead to increased uncertainty. Part of the uncertainty stems
from the perception that the safety of GMOs is still in question.
Another part stems web posts and media stories that may
inaccurately portray or overstate the risks and fail to tout the
benefits and the numerous consensus statements averring their
safety.
Similar trends have been observed in other developing
nations, even ones in the throes of a hunger crisis. Tens of
thousands of tonnes of corn donated by the United States was
rejected by Zambias president, who called it genetically mod-
ified poison. One stated rationale noted that since Europe
objects to its use, it must not be safe. While a large number of
Zambians are not welcoming to the idea of GMO, an increas-
ing fraction is warming (ever so slowly) to the possibility of
someday growing GMOs. However, someday sits far off on
the horizon as existing legislation all but bans the use of GM
seeds and proponents have yet to unify and work toward
acceptance.
A larger portion of Kenyans compared to those from other
nations in Africa are more accepting regarding the use of GMOs
partially due to wider acceptance by prominent political and
scientific leaders in Kenya. Greater acceptance and use of GM
crops are aided by two factors. First, the British left behind a
relatively advanced research center giving Kenyans in direct
contact with the latest scientific data on GMOs. Second, com-
mercial interests are exceptionally strong in Kenya, and they
promote the development and use of GM crops.
If Kenya is one of the more progressive nations regarding
54 FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer
have been grown in South Africa since 1997. As with Kenya,
South Africas educational system is built on deep scientific
roots. Local scientists affirming GMOs safety and benefits
help promote consumer acceptance of biotechnology. South
Africa, Egypt, Sudan, and Burkina Faso are the four nations in
Africa that sell GM crops commercially.
Latin America
Widespread production of GM crops in Brazil and Argentina
means that approximately one-third of the worlds acres
planted in GM crops occur in South America. GM maize
leads the production, but GM versions of cotton, soybean,
and canola are also grown in South America.
The widespread production of GM crops does not mean
South American consumers readily accept GM technology.
Interestingly, the countries with high production Brazil and
Argentina also have high resistance. Arguments against GMs
use include a lack of human resources and appropriate equip-
ment in the region, concerns about viability of small farmers,
disputes about seed ownership, and fears that regulation sur-
rounding GMOs is inadequate.
Peru and Colombia, in particular, are actively protesting
GMOs and resisting the efforts of multinational seed compa-
nies. Countries such as Mexico have actually banned the grow-
ing of GM crops. In 1998, it became illegal to grow GM maize
after the seeds were introduced into Mexico either through
accidental cross contamination or through illegal intent. How-
ever, the law has been loosely enforced. Reiteration of the ban
in 2013 by a judge may change this. Vocal leaders emphasize
corns essentiality and identity as Mexicos national crop and
describe GM corn as a threat to their sovereignty and identity.
Mexico faces opposition from such bodies as the United
Nations Food and Agriculture Organization, which argue in
favor of GMOs to eliminate hunger. Thus, across Latin Amer-
ica, mass scale production of GMO crops occurs while there are
bans and widespread resistance, sometimes within the bound-
aries of the same country.
GM Foods in Developing Countries: A Beacon of Hope?
In most developing countries, GM grains promise many agro-
nomic benefits. With the worldwide population expected to
double by 2050, food security is of great concern. Biotechnol-
ogy can be one of the most sustainable ways for ensuring
adequate agricultural yields and value-added and nutritionally
enhanced products. Statements such as those in the journal
GMO Biosafety Research dub the technology a beacon of hope.
Biotechnology protects food security via three primary
methods: increasing yields on existing land and optimizing
other limited resources, advancing food quality and safety,
and potentially bettering the economic status of local farmers.
GM grains can offer numerous nutritional advantages, such
as fortification to address nutrient gaps or extreme deficiencies
in developing countries. For example, WHO estimates that 250
million preschool children are vitamin A-deficient and an
estimated 250 000 to 500 000 vitamin A-deficient children
become blind every year, half of them dying within 12 months.
GM-produced golden rice adds vitamin A to help mitigate this
tragic public health problem.
No matter what the region, it is important that GM foods
and technologies address needs of local and subsistence
farmers, as well as respect cultural, dietary, and religious
importance of certain crops. In some cases, GM versions of
familiar crops may have greater acceptance than an unfamiliar
crop. While this is true in all regions, it may be especially true
for those in developing nations.
Judicious use of GM crops in developing nations means
that efforts must be directed to ensure that the use of this
technology does not exacerbate the economic divide. Care
must be exercised to ensure that the benefits of GM crops
help address the problem of food security and improve the
environment in the region and that economic or other risks are
minimized.
Conclusion
The practice of biotechnology began in the 1970s, and grains
such as maize were some of the first commercially available
GM foods. Biotechnology has been employed with grains to
heighten the ability to withstand stress and resist pests, disease,
and herbicides; enhance the nutrient profile, freshness, or
flavor of the plant; and reduce spoilage and improve yield.
Most crops available up until now have increased benefits to
farmers and this indirectly can help contain food costs for
consumers. Soon, foods with increased flavor and nutrient
profiles will be on the market.
Perception and acceptance of GM foods vary. The scientific
community concludes that GM foods, including GM grains, are
safe and pose no more risks than do conventionally grown
foods. Consumers remain wary despite the large body of sci-
entific evidence and numerous scientific and government bod-
ies deeming them safe and a possible way to address future
food needs. Attitudes toward GM foods strongly influence
perception of benefit and safety. Such attitudes are influenced
by media, legislation, science literacy, trust in regulatory orga-
nizations, price, religion and morality, and social economic
status.
It is exceptionally difficult to capture an entire populations
sentiment in one study, let alone a few sentences. Scientific
literature shows a lack of consensus, even within a region. It is
even more difficult to mention or represent every countrys
perspective. Despite limitations, this article divulges consumer
perception as it was most commonly portrayed in the scientific
literature. Overall, awareness of GM foods has increased sig-
nificantly worldwide. In the developing world, GM grains offer
the prospect of closing nutritional gaps, but still meet signifi-
cant resistance.
Many in the developing nations remain wary of biotech-
nology. Wariness is increased because most biotechnology
initiatives emanate from multinational companies. This fact
means that not only there may not be adequate understanding
of the consumers in a region but also there is worry about
motives of the multination and about who will reap economic
gains. Acceptance of biotechnology may increase when goals of
 FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer
developed countries align with those of developing countries,
and the benefits are made reciprocal and equal. In some
regions, the technology is slowly gaining and more hectares
are being planted.
The developed world is divided over the use of GM foods.
The United States, parts of South America, and South Africa
currently have large areas devoted to GM-crop production.
Australia and New Zealand place great weight on the benefits
and who reaps the benefits. In most regions, GM approval
continues to be higher in medicine than food, but GM food
approval still is relatively high in these regions. The most
vehement opposition comes from parts of Europe.
Where GM crops are allowed, many jurisdictions require
labeling and regulation of crops produced by biotechnology.
Mandated labeling on all GM foods and foods containing
GM ingredients is currently enforced in several countries in
Europe, Japan, Australia, Brazil, Russia, and China. In coun-
tries where no labeling of GM foods is required, labeling has
become a heated issue. Many consumers claim they have the
right to know what is in their food, and the government has
an obligation to make such information readily accessible.
Those opposed to labeling of GM foods cite consensus state-
ments of government agencies and health organizations stating
that food grown from GM seeds is not materially different
from food produced conventionally; therefore, there is nothing
to disclose. The US consumer population is largely in favor of
labeling, but federal labeling guidelines are voluntary. The
global status of labeling on GM foods may change in the
coming years, as consumers continue to demand transparency
from their governments. It may be interesting to observe how
consumer perception and purchasing behaviors of GM foods
are altered in response to increased transparency.
Encouragement of GM crops as a way to address the needs
of the world population in 2050 by humanitarians such as
those working for the Gates Foundation could have a dramatic
effect on attitudes and acceptance of GM foods in many parts
of the world. Further, open and transparent communication
on the benefits of GM foods and the availability of foods with
specific consumer benefits may help to address the concerns of
many consumers and improve acceptance.
Exercises for Revision
In one continent, take a sampling of countries from various
sectors and show in detail how these countries vary in their
acceptance and regulation of GMO.
Take a country or region and define the major concerns
regarding acceptance of GMO.
Take a country or region and suggest communication
strategies that would be convincing for the acceptance of
GMO.
What additional scientific information is needed for a cer-
tain population to accept GMO?
What world events or program backing has the potential to
impact GMO acceptance?
What problems has golden rice faced on its journey and
how could such things be avoided in the future?
55
Exercises for Readers to Explore the Topic Further
To what extent do countries have access to scientific litera-
ture concerning biotechnology and how does it affect their
overall consumer perception?
How well do GM grains address nutritional deficits in the
developing world?
How does labeling affect consumer perception of GM
foods?
See also: Food Grains and the Consumer: Labelling of Grain-
Based Foods.
Further Reading
American Medical Association, AMA (2013) Report of the Council on Science and
Public Health: Labeling of Bioengineered Foods. http://www.ama-assn.org/ama
(accessed 01.14.).
American Council for Science and Health (2014) The co-existence of organic and GMO
foods. http://acsh.org/2014/04/co-existence-organic-gmo-foods/ (accessed 4/2014).
Center for Food Safety (2013) US Polls on GE Food Labeling. http://www.
centerforfoodsafety.org/issues/976/ge-food-labeling/us-polls-on-ge-food-labeling
(accessed 01.14.).
DeSteur H, Gellynck X, Storozhenko S, et al. (2010) Willingness-to-accept and
purchase genetically modified rice with high folate content in Shanxi Province,
China. Appetite 54(1): 118125.
EuropaBio (2013) Abc stresses overwhelming weight of evidence on safety of GM
crops. http://www.europabio.org/news/abc-stresses-overwhelming-weight-
evidence-safety-gm-crops (accessed 01.14.).
European Commission (2006) Eurobarometer 66: Public Opinion in the European
Union. http://ec.europa.eu/public_opinion/archives/eb/eb66/eb66_highlights_en.
pdf (accessed 01.14.).
Hart J and Herrmann R (2009) Do they always say no? German consumers and second-
generation GM foods. Agricultural Economics 40(5): 551560.
Herath D (2001) Evolving Consumer Acceptance of Biotechnology Applications in
Canada: Evidence from the Public Opinion Surveys in 2001 (fifth wave) and 2011
(seventh wave). http://ageconsearch.umn.edu/bitstream/150525/2/Herath8_1AAEA.
pdf.
International Council for Science, ICSU (2003) New Genetics, Food, and Agriculture
Scientific Discoveries Societal Dilemmas. http://www.icsu.org/publications/
reports-and-reviews/new-genetics-food-and-agriculture-scientific-discoveries-
societal-dilemas-2003/ (accessed 01.14.).
International Food Information Council (2012) Consumer Perceptions of Food
Technology Survey. http://www.foodinsight.org/Content/5438/FINAL%20Executive
%20Summary%205-8-12.pdf.
International Service for the Administration of Agri-Biotech Applications, ISAAA (2013)
ISAAA Brief 442012: Press Release. http://www.isaaa.org/resources/publications/
briefs/44/pressrelease/ (accessed 01.14.).
Kaye-Blake W, Saunders C, and Fairweather J (2007) GM food and New Zealand
Consumers. http://www.biotechlearn.org.nz/themes/
new_zealand_views_on_biotech/gm_food_and_new_zealand_consumers
(accessed 01.14.).
Myhr AI (2007) The precautionary principle in GMO regulations. In: Traavik T and
Lim LC (eds.) Biosafety First Norway: Tapir Academic Publishers (accessed 01.14.).
http://genok.no/wp-content/uploads/2013/04/Chapter-29.pdf.
The National Academy of Sciences, NAS (2004) Safety of Genetically Engineered Foods:
Approaches to Assessing Unintended Health Effects. http://dels.nas.edu/Report/
Safety-Genetically-Engineered-Foods-Approaches/10977?bname (accessed
01.14.).
Nielsen T (2012) How do concerns about pesticides impact consumer willingness to
buy genetically modified french fries in Germany? Results from a purchasing
experiment. 2012 International European Forum, 1317 February, 2012. Austria:
Innsbruck-Igls.
Paulsen T GM food fight: Why the Gates Foundation wants to make rice golden. http://
www.humanosphere.org/basics/2013/09/gm-food-fight-why-the-gates-foundation-
wants-to-make-rice-golden/.
56 FOOD GRAINS AND THE CONSUMER | Genetically Modified Grains and the Consumer
Torres CS, Suva MM, Carpio LB, and Dagli WB (2006) Public Understanding and
Perception of the Attitude Towards Agricultural Biotechnology in the Philippines.
ISAAA. SEARCA and CDC-UPLB. http://www.isaaa.org/programs/supportprojects/
(accessed 01.14.).
USDA, Economic Research Service (2013) Adoption of Genetically Engineered Crops in
the U.S. www.ers.usda.gov/data-products/adoption-of-genetically-engineered-
crops-in-the-us/recent-trends-in-ge-adoption.aspx#.U0FCRMf95lk (accessed
01.14).
Vinter R (2013) Livestock 2013: Survey shows consumer resistance to GMs. Farmers
Weekly. http://www.fwi.co.uk/articles/03/07/2013/139866/livestock-2013-survey-
shows-consumer-resistance-to.htm (accessed 01.14.).
World Health Organization, WHO (2014) 20 questions on genetically modified foods.
http://www.who.int/foodsafety/publications/biotech/20questions/en/ (accessed
01.14.).
Zhang X, Huang J, Qiu H, and Huang Z (2010) A consumer segmentation study
with regards to genetically modified food in urban China. Food Policy 35(5):
456462.
Relevant Websites
http://www.agbioworld.org/ AgBio World, Peer Reviewed Publications on the Safety of
GM Foods.
http://dels.nas.edu/ National Academy of Sciences, Division on Earth and Life
Studies. National Research Council (U.S.).
http://edis.ifas.ufl.edu/fe934 Evans, EA Ballen, FH. A synopsis of US consumer
perception of genetically modified (biotech) crops. University of Florida Extension.
http://www.ers.usda.gov/data-products/adoption-of-genetically-engineered-crops-
inthe-us/ Fernandez-Cornejo J, 2011 U.S. Economic Research Service, Adoption
of Genetically Engineered Crops in the U.S.
http://www.gmo-compass.org/eng/agri_biotechnology/gmo_planting/ GMO
Compass.
http://www.icsu.org/ International Council for Science.
http://www.isaaa.org/ International Service for the Acquisition of Agri-biotec.
 Labelling of Grain-Based Foods
JM Jones, St. Catherine University, St. Paul, MN, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Basic label requirements are similar across jurisdictions
with the ingredient statement and contact information
being critically important.
Label terminology used to describe grain processing,
milling, and grain fractions is understood by industry, but
not by consumers. Further, it is not standardized across
jurisdictions and is confusing to consumers.
Nutrition labeling of grain-based foods is mandatory in
many countries. Whether required or voluntary, the format
is specified but formats vary across jurisdictions.
Regulation for labeling of organic and genetically modified
grains and other label descriptors such as fresh and natu-
ral vary across jurisdictions.
Procedures and rules regarding the approval and use of
nutrient-content claims, structurefunction claims, and
health claims in grain-based foods depend on the
jurisdiction.
Issues regarding the labeling of whole-grain foods are com-
plicated due to varying recommendations for whole-grain
intake across regions, issues about the amount of dietary
fiber or detractor nutrients that are in the whole-grain food,
and the use of wet or dry weight for calculation of the whole
grain.
Learning Objectives
To be able to outline and define various requirements and
terms associated with labeling grain-based foods.
To compare various label descriptors, nutrition labels, and
types of nutritional claims used on grain-based foods in
various jurisdictions.
To delineate issues associated with labeling of whole grains
and whole-grain foods within and across jurisdictions.
Overview
Grain-based food (GBF) labels allow consumers to select and
purchase products with desired characteristics, ingredients, and
nutritional profiles. Labels act as both marketing tools and
transmitter of legally required information. They must name
and accurately describe the food and list the ingredients in
descending order by weight. The ingredient statement is critical
for most consumers as it delineates what is in the product and
gives the consumer an idea of the relative amounts of the
various components in a product, since they must be listed in
order by weight from the most to the least prevalent. In some
countries, percent labeling is required, so that a precise amount
of a specific ingredient is declared. Interested consumers can
carefully scrutinize the ingredient statement for components or
additives that they wish to include or avoid in their diets.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00076-0
Coded or overt dates and lot numbers are required both to
track shelf life and to allow for product recalls. The manufac-
turer contact information enables consumers to make queries
and to state complaints and facilitate recalls, if necessary. The
net weight allows price and value comparisons of competing
products and ensures the consumer receives a fair measure.
Some jurisdictions may require the country of origin labeling,
warning statements about allergens or ingredients of concern,
and advisory statements on safe handling of the product.
Nutrition labeling, usually with a prescribed format, is
mandatory in many jurisdictions, while others allow or
encourage it voluntarily. Statements and visuals on the package
and advertising must not be misleading. Foods for special
dietary or medical use such as cereals or bars that have func-
tional ingredients that reduce blood sugar may have more
extensive nutrition labeling requirements. Nutrient-content,
structurefunction, and health claims are allowed in some
jurisdictions, but specific claims and rules regarding them
vary across jurisdictions.
GBFs also carry label descriptors about the size, shape, and
processing of the grain. In many cases, the terminology is
familiar only to those in the cereal industry. Thus, consumers
are confused about definitions surrounding grain fractions,
whole grain (WG), and especially the definition of a whole-
grain food (WGF). This article will discuss these and other
issues specific to GBF.
Label Descriptors for GBF
Label descriptors for GBF can be those that are more general
such as fresh and those that communicate aspects regarding
the product, form, and level and type of processing specific to
grain such as the method and degree of milling, the particle
size, and other processing aspects. While these industry terms
associated with GBF are understood by industry, they can leave
consumers confused. Terms such as medium pearled or de-
germed are not always as clear or as easy to operationalize as
terms for other common foods, such as canned sliced peaches
in syrup, where most of these terms need little further clarifi-
cation. Nonetheless, this information is needed for selecting
the grain form for the best culinary use and for choosing
desired nutritional characteristics.
Terms such as fresh, natural, homemade, artisan, tra-
ditional, ancient, and stone-ground are popular with mar-
keters and consumers alike but have come under scrutiny by
various regulatory bodies. For the consumer, such terms give
products a goodness halo that may or may not be justified, so
some jurisdictions have prescribed ways such terms can be
used. In areas where there is no legal definition, the expecta-
tion is that such terms are not misleading. Terms such as
natural or fresh may or may not be defined depending on
the jurisdiction. In some jurisdictions, the term fresh for bread
is defined to mean prepared at the venue and sold there
57
58 FOOD GRAINS AND THE CONSUMER | Labelling of Grain-Based Foods
shortly after baking. Preprepared dough off-site and baked on
the premises cannot be labeled as fresh in some regions. It is
easy to understand why terms are confusing because the term
fresh for pasta denotes that the pasta is not dried.
Designations such as light, low-calorie, very low sodium, Rules about saturated fat, trans fat, and calories from fat vary
high in, or free from may be strictly defined or not allowed.
For example, in the United States, rice cakes with under 40 cal
might be labeled as a naturally low-calorie food and cannot
be labeled as low-calorie rice cakes implying that they are
lower in calories than other rice cakes. In certain jurisdictions,
a GBF formulated to reduce an attribute such as calories,
sodium, or fat must be reduced by a specific percent and
must be labeled and compared to either the original food or
the average of foods in the category.
Claims that a food is high in x may need to follow rules for
nutrient-content claims and may not be allowed for compo-
nents that are not nutrients such as WG or antioxidants. In
some jurisdictions, free from labeling is specified in that foods
that have never had component x cannot be labeled as free
from x. Statements about gluten-free must follow rules of the
jurisdiction. The CODEX regulations, followed in a number of
areas, require < 20 ppm gluten, but a few regions also allow
low gluten with higher levels of gluten. Allergen labeling may
be required, and so GBF would have the required disclosures of
wheat, nuts, milk, eggs, and other common allergens.
Organic labeling for GBF aids consumers wishing to purchase
products that are produced without additives, pesticides (other
than those allowed by organic rules), and chemical fertilizers
and did not undergo genetic modification (GM) or irradiation.
Some regions require official certification at all stages of the
process in order to label products as organic. Table 1 shows a
gradation in organic that is allowed in the United States with
products where 95% or more of the product is organic can bear
an official seal. If a product has < 70% organic ingredients, it
cannot use organic on the main package label, but individual
ingredients listed as organic on the ingredient statement.
If GM grains are allowed, many countries require their
labeling. In the United States, labeling of GM grains is not
required because the USFDA views that there is neither a mate-
rial difference nor a safety or environmental issue associated
with these products. However, deeply held beliefs by some
consumers have prompted a number of state legislators to
sponsor bills requiring GM labeling. To date, all such initia-
tives have failed.
The Nutrition Label
The nutrition labeling of packaged foods, including GBF, is
mandatory in many countries of the world. Where voluntary,
Table 1 The USDA organic label definitions
100% organic Must contain only organically produced
ingredients
Organic At least 95% of the products ingredients
must be organic
Made with organic Must contain at least 70% organic
ingredients ingredients
Some organic Products contain <70% organic ingredients
ingredients
most countries have regulated formats. The nutrition label in
most countries requires a declaration of energy in calories or
joules, total fat, saturated fat, trans fat, cholesterol, sodium,
total carbohydrates, dietary fiber (DF), sugars, and protein.
across jurisdictions. Similarly, the list of micronutrients
required and allowed varies as does the percent daily value
(DV) standard. In most jurisdictions, labeling becomes com-
pulsory if a nutrition claim regarding a specific nutrient is on
the label or in advertising. Nutrition labeling is harmonized
throughout the European Union and some regions; however,
this is often not the case.
DF values in GBF may vary depending on which DF defini-
tion and analytic method have been officially accepted. The
acceptance of Codex Alimentarius and aligned DF definitions
and methods by Europe, Canada, and many other jurisdictions
has helped harmonize fiber labeling across many jurisdictions.
Energy values may also vary depending on what calorie value is
assigned to certain fibers in various regions.
In most countries, nutrition labels are based on 100 g of
food, but some countries use dual labeling. Nutrient values are
listed per 100 g of food and also for a serving size appropriate
to the food and the country. In the United States, the nutrient
values are based only on government-mandated serving sizes
called reference amounts commonly consumed (RACCs).
The system using 100 g portions has the advantage of allow-
ing easy comparison of products and enables consumers to
easily calculate the percent of various macronutrients. How-
ever, some consumers cannot envision how 100 g of a specific
product a serving can be eaten. The following examples illus-
trate this confusion. A slice of bread is usually 30 g so labeling
per 100 g would be the nutrient value for three slices of bread.
Ready-to-eat cereals vary dramatically so a 30 g serving of a
dense granola is 1/3c, whereas it is two cups of a puffed grain
cereal. Unfortunately, many consumers are not able to use the
nutrition label effectively as they are unable to apply the listed
values to the amount they have eaten.
The nutrition label must list the nutrients in the package as
sold, regardless of how the food is commonly ingested. Thus,
some products have two values the one required for the food
as packaged and the one that gives nutrition information of the
product as prepared or ingested. Thus, cereal labels often give
values for the cereal as purchased and as consumed with 120 g
(c) of low-fat milk. A similar type of dual labeling may be on
packaged foods such as cake or pasta mixes, which have an
additional column giving nutrient values for the food prepared
according to package directions. Dual-language (or multilan-
guage) labeling of GBF enables their sale in countries such as
Canada with more than one national language or in a region
such as Southeast Asia.
Labelling of the Various Grain Components
Grains have been processed since before biblical times to
remove the outer inedible husk, and in many cases, outer layers
may be removed. Grains may then be broken into particles of
various sizes, sifted into various streams, and kept separate or
recombined, and the various fractions may be processed fur-
ther. The specific terminology used to describe grain
 FOOD GRAINS AND THE CONSUMER | Labelling of Grain-Based Foods
components from various stages of milling and processing is
on the label, but consumers do not know if a meal is different
from a grit and may think that degermed is a sterilization, and
not a milling, process.
In many parts the world, GBFs are most commonly made
from refined or enriched grains. These foods have mostly the
endosperm of the grain, or in the case of enriched, the endo-
sperm has some nutrients lost in milling added back. Current
nutrition research suggests that WGs are advantageous in terms
of more DF and reduced risk of chronic disease. Thus, some
consumers are trying to replace refined and enriched grain
foods with WGFs, which makes the understanding of GBF
label terms so important.
Enriched and Refined Grains: If the various parts of a kernel
have been removed, it is a refined grain. If some vitamins and
minerals lost in milling are added back, the grain is enriched.
Various food authorities decide if enrichment is allowed and, if
so, the specific nutrients added and the amount allowed. If
enriched flour is used in packaged food products, the nutrients
added not only must be reflected on the nutrition facts panel
but also may be reflected on the ingredient statement. The
ingredient statement might read Ingredients: Sugar, Enriched
Bleached Flour (wheat flour, niacin, iron, thiamine mononi-
trate, riboflavin, folic acid). . ..
Grains may also be fortified to address nutritional inade-
quacies in the region. Thus, the nutrient levels of fortified
products may be higher than enriched products. Often, 25%
of the DV is added, but some products have more. These
additions are reflected on both the ingredient statement and
the nutrition facts panel. If fortification levels in some GBF
meet the daily requirement, for example, 100% of the DV, the
product must be labeled as a multivitamin and mineral sup-
plement GBF. The intake of these fortified cereal products
increases intakes of nutrients of concern. In the case of folic
acid fortification of grains, there has been a reduction of
neural tube defects, and certain birth defects have been
decreased by 5070% in various countries. Yet some fear
that such a policy could put some groups at risk for excess
folic acid intake, even though most data do not indicate that
this might be a problem.
WG was first defined by the American Association of Cereal
Chemists (now AACCI) in 1999 as follows:
Whole grains shall consist of the intact, ground, cracked or flaked
caryopsis, whose principal anatomical components the starchy
endosperm, germ and bran are present in the same relative pro-
portions as they exist in the intact caryopsis.
WGs have subsequently been defined by other organiza-
tions. The Whole Grains Council (WGC) created a more
consumer-friendly definition. It reads as follows:
Whole grains or foods made from them contain all the essential
parts and naturally-occurring nutrients of the entire grain seed. If the
grain has been processed (e.g., cracked, crushed, rolled, extruded,
lightly pearled and/or cooked), the food product should deliver
approximately the same rich balance of nutrients that is found in
the original grain seed.
The European HEALTHGRAIN (HG) workshop (2008)
defined WG as follows:
59
Wholegrain is defined as intact and/or processed (e.g., de-hulled,
cleaned, ground, cracked, flaked or the like) grains, where the frac-
tions endosperm, bran and germ are present in the same proportion
as found in the least processed traditional forms of the edible grain
kernel of the same species.
All three definitions do not require that the grain remain as
an intact kernel, but rather that all of its components be pre-
sent. However, a few countries using the terminology require
that whole defines the intact kernel. Each definition varies
slightly in terms of both consumer friendliness and allowable
loss of outer layers. The WGC definition is more consumer-
friendly than the AACCI definition that uses the botanical term
caryopsis. The HG definition allows some removal of a small
amount of grain such as that occurs in traditional processes
such as making bulgur or nixtimalizing corn or those that
make the grain healthier. Examples include the scoring of the
outer kernel to increase availability of nutrients or nixtimaliz-
ing to reduce mycotoxins in the outer layers. Some countries
equate whole-wheat or wholemeal flour; others allow the
removal of some of the outer layers and still call it whole-
wheat flour. This does create confusion.
Definitions of WG help processors formulate food and help
consumers describe WGFs that are all WG such as porridge oats
(oatmeal) or WG flours. However, the consumer has great diffi-
culty determining which foods are WGF or determining the num-
ber of servings of WG to ingest in products that are partly WG.
Each current strategy for finding WGFs in the marketplace
has a disadvantage. One strategy suggests that a food is WG, if
the first ingredient in the ingredient statement is WG. While
this may correctly identify a WGF, it may not if the food has
many ingredients. This is explained by the following examples.
The ingredient statement from a bread mix is as follows:
enriched flour, whole-wheat flour, brown rice flour, rolled
oats, nonfat dry milk, yeast, and salt. This could be a bread
mix with 90% enriched flour and 10% other ingredients mak-
ing it not a WG bread mix. On the other hand, the ingredient
statement could reflect 35% refined flour, 30% whole-wheat
flour, 30% brown rice flour, 3% rolled oats, and 2% other
ingredients. This bread mix would clearly be a WGF even
though refined flour was the first ingredient. A 30 g serving of
granola could have 5 g of whole oats; 4 g each of peanuts,
almonds, pecans, and walnuts; and 5 g of raisin paste, honey,
and salt. The granola, while contributing WG and many nutri-
tious ingredients, may not give a dietarily significant amount of
WG. Some suggest that combining the ingredient statement
with the products DF levels could identify WGFs, but this
approach works only if the formulation contains high-fiber
WGs. Low-fiber WGs such as brown rice would have a low-
fiber level on the nutrition panel.
Achieving international agreement on the amount of WG
that should be in a product to call it a WG serving is very
difficult. There is no international agreement regarding the
recommended daily intake of WG. Some regions make no
recommendations. For those that do, recommended intakes
vary from 48 to 115 g day 1. This wide range makes agreement
on the amount of WG in a product to call it a WGF-difficult.
Some jurisdictions allow manufacturers to label statements
of fact, which means statements such as YXZ product gives xx
grams of WG per serving. The difficulty is that most consumers
60 FOOD GRAINS AND THE CONSUMER | Labelling of Grain-Based Foods
do not know the amount of WG recommended per day, so such
labeling could easily be misinterpreted. Similarly labeled ban-
ners such as made with whole grain, whole-grain goodness, detractors like sugar or fat in the product. For some consumers,
and multigrain and even high-fiber make some consumers any designation of a food as WG is misinterpreted and creates a
think that the food meets one of the recommended WG servings.
The WGC stamp has high consumer recognition, is easy to
understand, and is used internationally. The WGC has a stamp
on the food label that indicates the product has 8 g of more of
WG per labeled serving or RACC. The stamp has two disadvan-
tages. First, it causes some consumers to deduce that only those
products bearing the stamp are WG. This may occur because
consumers are unaware that some companies may not use the
stamp because of its cost. Its second disadvantage is that the
stamp gives an unfair advantage to products with low moisture
or large serving sizes. This creates a scenario where a dough
made into a 25 g bun would not have the needed 8 g of WG to
receive the stamp, but the same dough in a 55 g bagel would.
Since a clear WG characterization was suggested as being
important to the goal of eating more WG, AACCI and HG tried
to respond by putting forward a way to identify WGFs. The
following questions were laboriously considered in the WGF
determination:
1. Should the WG characterization apply only to products that
are in the grain group?
2. Should it be based on a serving of food or be proportional
and therefore serving size-neutral?
3. Should it be based on a wet (as served) or dry weight basis?
(Use of wet weight favors dry products and penalizes
cooked oatmeal, bread, and soup with pasta or barley.)
4. Should it apply only to the grain portion of the food? If so,
should there be a minimum level or grain, even if all the
grain is WG?
5. Should it indicate that the food would deliver a certain
amount of DF or not have a certain level of detractor
nutrients such as sugar, salt, fat, or saturated fat?
6. Should it apply to the entire meal in a frozen or other
dinner where the side of grain would meet the requirements
but would fail to do so when all the food entree and other
meal components are factored into the calculation?
7. Should half the grain in the product be WG?
The AACCI Whole Grain Working Group put forward a WG
characterization that suggests a proportionality of 8 g of WG
per 30 g of food. This addresses problems created by small and
large serving sizes, but leaves some unanswered questions such
as WG pasta in a soup or side in a ready-to-eat meal. The 8 g/
30 g does not specify that the product must have half of the
grain be WG. Rather, it places a WG minimum, which allows
formulators of GBF to increase the other grain fractions such as
bran, germ, or aleurone, which technically are not WG but
carry many nutritional attributes of WG. Some are concerned
that the characterization could allow products to have higher
levels of refined grain component than WG. This could be true
but the intent of using the 8 g per 30 g is that with the recom-
mendation of six 30 g servings of GBF needed for most calorie
levels, a person would get a minimum of 48 g of WG, an
amount thought to be dietarily important.
It should be noted that it is merely a characterization and
does not speak of the nutritional attributes such as DF or
halo effect causing consumers to think such foods have more
DF, fewer calories, and detractor ingredients than they do.
Further, this characterization is only voluntary and the net
result is that there can be many package statements claiming
WG content and consumer confusion.
Nutrient Content, Health, and StructureFunction
Claims
Nutrient-content claims are allowed in some jurisdictions pro-
vided that they are factual and use terminology in ways spec-
ified in the regulations. In many cases, a serving of a food
carrying a nutrient-content claim must provide a certain min-
imum percentage of the DV, often 10% in a normal serving.
For example, Acme cereal is a significant source of folic acid.
Structurefunction claims are popular in some countries.
Japan was the first to enfranchise these and allow a number
such claims. Such claims are also allowed in North America,
Europe, Australia, and parts of Asia. Structurefunction claims
may point out the association with health but must not claim
to cure a disease. Claims such as Acme cereal is a significant
source of vitamin A, which has been associated with skin and
eye health, and Bran fiber in Acme cereal helps maintain
bowel regularity are allowed in some jurisdictions. As with
structurefunction claims, the amount of nutrient needed to
make a nutrient-content claim is often the same. Some coun-
tries prohibit any claims if the food also contains the so-called
detractor nutrients, such as saturated fat. If a claim is allowed,
the detractor nutrient must also be prominently labeled in the
same place as the positive claim. In order to have claims on a
food that diagnose, mitigate, treat, cure, or prevent a disease or
specific class of diseases, officially approved health claims are
used. Health claims are accepted only after significant scientific
agreement by the regulatory authorities in the region. The only
exception is that of classical nutrient deficiency diseases.
Table 2 gives a sampling of health claims for GBF.
Health Claims Regarding Blood Glucose
The Swedish Nutrition Foundation (SNF) allows the following
claim about blood glucose: oat bran -glucan reduces blood
glucose and insulin response.
EFSA allows a claim on a lowered glucose response if 14 g of
resistant starch is substituted for 14 g of available starch.
Summary
Labeling of GBF informs consumers when making grain pur-
chases, enables industry to comply with rules regarding GBF
and grain components, and helps regulators ensure consumer
 FOOD GRAINS AND THE CONSUMER | Labelling of Grain-Based Foods
Table 2 Health claims for grain-based foods in various jurisdictions
Fiber-containing grain products, fruits, and vegetables and cancer
Sample claim: Low-fat diets rich in fiber such as Acme Bread may reduce
the risk of some types of cancer, a disease associated with many
factors
Requirements: GBF must meet criteria for low fat and, without
fortification, be a good source of dietary fiber
Fruits, vegetables, and grain products that contain fiber, particularly
soluble fiber, and the risk of coronary heart disease
Sample claim: Diets low in saturated fat and cholesterol and rich in fruits,
vegetables, and grain products that contain some types of dietary fiber,
particularly soluble fiber, may reduce the risk of heart disease, a
disease associated with many factors
Requirements: GBF must meet criteria for low saturated fat, low fat,
and low cholesterol. They must contain, without fortification, at least
0.6 g of soluble fiber per reference amount, and the soluble fiber
content must be listed
Folic acid and neural tube defects
Sample claim: Healthful diets with adequate folic acid may reduce a
womans risk of having a child with a brain or spinal cord birth defect
Requirements: Enriched GBF must meet or exceed criteria for good
source of folic acid, that is, at least 40 mg of folic acid per serving (at
least 10% of the daily value)
Dietary soluble fiber, such as that found in whole oats, barley, oat bran,
and Oatrim or psyllium seed husk and coronary heart disease and
cholesterol lowering
Sample claim: Diets low in saturated fat and cholesterol that include 3 g
of soluble fiber from whole oats per day may reduce the risk of heart
disease. One serving of this whole oat product provides xx g of this
soluble fiber
Requirements: Foods must meet criteria for low saturated fat, low
cholesterol, and low fat. Foods that contain whole oats must contain
at least 0.75 g of soluble fiber per serving. Foods that contain psyllium
seed husk must contain at least 1.7 g of soluble fiber per serving
Typical foods: GBF made with whole or rolled oats, oat bran, or whole oat
flour; barley; psyllium seed husk; and isolated or modified b-glucans
from barley and oats
Note: The European Food Safety Authority (EFSA) allows the claim Oat
beta-glucan has been shown to lower/reduce blood cholesterol. Blood
cholesterol lowering may reduce the risk of heart disease. The Swedish
Code of Practice allows Soluble fiber from oat bran may help reduce
cholesterol. The United Kingdom allows The inclusion of oat bran can
help reduce cholesterol as part of a diet low in saturated fat and a
healthy lifestyle. The Netherlands Nutrition Centre (Voedingscentrum)
allows One daily portion of ACME oat bran-based bread reduces
cholesterol within 3 weeks and helps maintain healthy arteries and
blood vessels. And Switzerland BAG Oat bran favorably impacts
cholesterol levels
Whole grain and heart disease
Diets rich in whole-grain foods and other plant foods and low in total fat,
saturated fat, and cholesterol may reduce the risk of heart disease and
some cancers
Dietary fiber content of at least 1.7 g per 35 g serving
Typical foods: Whole-grain breads and cereals (USFDA claim)
WG claims not been approved in many other jurisdictions because of
inadequate characterization of the ingredients and rulings of inadequate
evidence
Vitamins B12, B6, and folic acid and reduced risk of heart disease
Sample claim: Foods containing vitamin B12, as well as B6 and folic acid,
may reduce the risk of coronary heart disease
Typical foods. Fortified GBF may make such claims in the United States
61
protection and fair marketing of GBF. Understanding label
terms for any food is challenging, but terms for GBF, which
describe grain forms, particle size, degree of milling and spe-
cific milling and processing treatments of various grain types,
and fractions, add to consumers confusion. This is com-
pounded because terms vary across regions and among grains
for a similar process. Terms such as enriched and refined are
applied to grains and not to most other foods. General label
terms such as fresh and organic apply to GBF, but these may
or may not be defined in a jurisdiction. If defined, definition
even within a jurisdiction may vary among products with
fresh on bread meaning something different than fresh on
pasta. Some use organic labeling as a way to avoid chemicals
and GMFs. Nutritionally labeled products are found in many
regions and help consumers meet individualized nutritional
needs, although many consumers are confused by many terms
on the nutrition and grain label. GBF can carry a number of
nutrient-content, structurefunction, and health claims. Each
region has specific rules regarding their approval and use.
Various health promotion agencies are suggesting the inges-
tion of more WG. However, the ability of consumers to make
informed choices as to what is a WGF is difficult. Even experts
disagree on the best and fairest strategy for such labeling. Vari-
ous strategies have been proposed for the identification of label-
ing WGFs, but each has some drawbacks. Most recently, AACCI
has put forward a WG characterization, which is a proportional
minimum of 8 g or WG/30 g of product. Labeling of GBF is
important and complicated. It varies across regions due to non-
standardized use of terms, varying nutritional requirements,
differing regulations by food authorities, and varying degrees
of consumer understanding, needs, and perceptions.
Exercises for Revision
Take a country or region and compare the nutrition label
and general labeling requirements for GBF.
Compare the various terms used to describe the milling
process of various grains such as pearling, polishing, pin
milling, and rolling and resultant particulate fractions such
as kernels, groats, grits, meals, and flours.
Compare health claim approval and use for GBF in various
countries.
Compare the labeling of WGs and WGFs across
jurisdictions.
Compare the rules for using the terms natural, fresh, and
organic across jurisdictions.
What contributions do enriched, refined, fortified, and
WGs make to the diet of a particular country or
subpopulation?
Exercises for Readers to Explore the Topic Further
Label marketing statements, grain and processing
terminology, and nutrition labeling are confusing to con-
sumers. Suggest some strategies to diminish confusion.
62 FOOD GRAINS AND THE CONSUMER | Labelling of Grain-Based Foods
What activities and labeling initiatives can be undertaken to
help consumers identify WGFs and increase their WG
intake?
Percent ingredient labeling is used in some countries for all
foods including GBF. Does this improve consumer under-
standing of what is in the food?
See also: Food Grains and the Consumer: Fortification of Grain-
Based Foods; Genetically Modified Grains and the Consumer; Grains
and Health: Misinformation and Misconceptions; Scientific
Organizations Related to Grains: Research Organizations of the
World: Global Trends and the Commercial Sector; Scientific Societies
Associated with Grain Science; The Basics: Grain: Morphology of
Internal Structure; The Cereal Grains: Barley: An Overview of a
Versatile Cereal Grain with Many Food and Feed Uses; Maize:
Overview; Oats: Overview; Rice: Overview; Wheat: An Overview of the
Grain That Provides Our Daily Bread.
Further Reading
AACCIs Whole Grains Working Group Unveils New Whole Grain Products
Characterization (2013). http://www.aaccnet.org/about/newsreleases/Pages/
WholeGrainProductCharacterization.aspx.
Barr SI, DiFrancesco L, and Fulgoni 3rd VL 3rd (2013) Consumption of breakfast and
the type of breakfast consumed are positively associated with nutrient intakes and
adequacy of Canadian adults. Journal of Nutrition 143(1): 8692.
Berry RJ, Bailey L, Mulinare J, Bower C, and Folic Acid Working Group (2010)
Fortification of flour with folic acid. Food and Nutrition Bulletin 31(Supplement 1):
S22S35.
FDA Wants Consumers to Truly Get the Whole Grain (2014). http://usgovinfo.about.
com/od/consumerawareness/a/wholegrain.htm.
Grieger JA and Cobiac L (2012) Comparison of dietary intakes according to
breakfast choice in Australian boys. European Journal of Clinical Nutrition 66(6):
667672.
Hamner HC, Tinker SC, Berry RJ, and Mulinare J (2013) Modeling fortification of corn
masa flour with folic acid: The potential impact on exceeding the tolerable upper
intake level for folic acid, NHANES 20012008. Food and Nutrition Research
57, 19146.
Holmes BA, Kaffa N, Campbell K, and Sanders TA (2012) The contribution of breakfast
cereals to the nutritional intake of the materially deprived UK population. European
Journal of Clinical Nutrition 66(1): 1017.
Yang Q, Cogswell ME, Hamner HC, et al. (2010) Folic acid source, usual intake, and
folate and vitamin B-12 status in US adults: National Health and Nutrition
Examination Survey (NHANES) 20032006. American Journal of Clinical Nutrition
91(1): 6472.
Relevant Websites
http://wholegrainscouncil.org/whole-grains-101/definition-of-whole-grains
HealthGrain Forum. Definition and analysis of whole grain.
http://wholegrainscouncil.org/whole-grains-101/definition-of-whole-grains Whole
Grains Council. Definition of whole grains.
 Grains and Health
R Korczak, University of Minnesota, St. Paul, MN, USA
D Hauge and B Maschoff, Grains for Health Foundation, St. Louis Park, MN, USA
L Marquart, University of Minnesota, St. Paul, MN, USA; Grains for Health Foundation, St. Louis Park, MN, USA
P Jacques, Tufts University, Boston, MA, USA
R Lindberg, Minneapolis Heart Institute Foundation, Minneapolis, MN, USA
R Menon, The General Mills Bell Institute of Health and Nutrition, Minneapolis, MN, USA
2016 Elsevier Ltd. All rights reserved.
Abbreviations
AFH Away from home
CBPR Community-based participatory research
CDC Centers for Disease Control and Prevention
PHAD Practical, healthier, affordable, and desirable
Topic Highlights
Innovative approaches are necessary to address the complex
and dynamic health-related issues facing food scientists,
nutrition and health, and community involvement.
Systems thinking is one approach to view problems as
parts of an overall system, rather than reacting to a specific
part, outcome, or event.
For example, how will we address the complexity of what
and how to deliver healthy grain-based foods that con-
sumers find acceptable and will include in their daily
diet?
At minimum, public health officials are being asked to
consider not only the overall roles of each player within
the food system and how they work together to create the
food environment but also the significance of community
and citizen representation in identifying and filling the
gaps, such as research projects that help meet the diverse
and complex demands of the public.
In order to increase consumer intake of these grain-based
foods with higher whole grain content, it will require the
development of tools, methods and approaches, systems,
and necessary infrastructure to successfully develop and
deliver these foods into the marketplace.
Public and private partnerships can create a collaborative
environment that maximize cross-disciplinary expertise
among government, academic, and industry researchers to
help solve complex research problems.
Food and ingredient industries are uniquely positioned
with the technical expertise to make gradual product
formulation changes that are well accepted by consumers.
Learning Objectives
Among the few major learning outcomes expected from
studying this article, the reader will be able to
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00067-X
PPP Public and private partnerships
USDA-ARS United States Department of Agriculture
Agricultural Research Service
WG Whole grain
identify challenges facing current research approaches used
to investigate the efficacy and use of grain-based foods by
consumers;
understand systems thinking for problem identification
and use collective solutions for increasing healthier grain-
based foods in the marketplace;
discuss the pros and cons of public and private partnerships
for advancing grain-based research;
describe the process of undertaking public and private
partnerships from (1) achieving a common purpose
among individuals, disciplines, and sectors; (2) sharing of
knowledge in a safe environment; and (3) arriving at
collective solutions;
outline one project that meets the public and private
partnership criteria related to grain-based foods issue.
Introduction
This article provides one perspective about our initial
approaches to address research initiatives relative to grain-
based foods that might benefit both industry interests and
nutrition-related public health issues. Research is proposed to
be accomplished through public and private partnerships
(PPPs) where the government and industry collaborate and
work toward collective solutions while comingling funding
sources from public and private sectors. Systems thinking
approaches and public nutrition policy will be addressed rela-
tive to identifying research initiatives that fill crucial gaps in the
supply chain, food environment, and local community. Three
proposed grain-based projects will be presented with rationale
for potential support through public and private research
efforts. Further discussion will focus on the appropriateness
of these projects as opportunities for PPP research initiatives.
This article is based on the initial steps leading to a 35-member
PPP research meeting focusing on healthier grain-based foods
December 2014 in Washington, DC.
63
64 FOOD GRAINS AND THE CONSUMER | Grains and Health
Moving from Traditional Research and Analytic
Thinking to Include Systems Approaches
Traditional nutrition research, from a biological and physio-
logical dimension, has been primarily explored through
in vitro, animal, observational, and experimental approaches
where reductionism is considered the gold standard. Analytic
thinking has been the mainstay for scientific investigation and
remains an important aspect of our current scientific practice.
These traditional approaches were extremely effective in
addressing nutrient deficiencies in the late nineteenth and
early twentieth centuries, up until the 1950s and 60s when
chronic diseases became a major issue of concern for food
and nutrition researchers.
As our society moves into a new millennium, innovative
approaches are necessary to address the complex and dynamic
health-related issues facing scientists through food technology,
nutrition and health, and community involvement. Unfortu-
nately, we are routinely taught throughout our academic
careers and professional lives on how to solve problems, but
rarely taught how to identify problems. According to Russell
Ackoff, We fail more often not because we fail to solve the
problems we face, but because we fail to face the right
problem. For example, how will we address the complexity of
what and how to deliver healthy grain-based foods that con-
sumers find acceptable and will include in their daily diet? One
solution, as recommended by the 2010 Dietary Guidelines for
Americans Committee, is for the sectors of government, industry,
and nonprofits (including both large and small businesses) to
come together in order to develop a system that would allow for
health-promoting dietary patterns. At minimum, public health
officials are being asked to consider not only the overall roles of
each player within the food system and how they work together
to create the food environment but also the significance of
community and citizen representation in identifying and filling
the gaps, such as research projects that help meet the diverse and
complex demands of the public.
For example, one way to fulfill the gap(s) and meet the
demands of the public is to develop a grain-based food that
is practical, healthier, affordable, and desirable (PHAD) such
that consumers will eat it in a restaurant setting. Yet the food
must be practical for supply chain stakeholders along with
chefs and clientele in restaurants. A grain-based food might be
described as healthier, such that whole grain (WG) wheat is
incrementally or gradually increased if the food/product
meets PHAD criteria and restaurant clientele actually eat it.
Affordability throughout the supply chain and in the food
preparation and restaurant setting can be a limiting factor.
Overall desirability of these grain foods by supply chain
stakeholders, restaurant chefs, staff, and clientele along with
community norms needs to be taken into account.
In order to increase consumer intake of these grain-based
foods with higher WG content, it will require the development
of tools, methods and approaches, systems, and necessary
infrastructure to successfully develop and deliver these foods
into the marketplace. The successful outcome of this effort
might be underpinned via research initiatives that focus on
increasing WG ingredients in grain-based foods for restaurant
service. Ideally, these inherent research problems and solutions
are conceived and tested within the context of a complex grain
foods supply, which interfaces with a community of clients
who frequent the food environment, such as restaurants
whereby these healthier grain-based foods are served and
potentially consumed.
Systems thinking is one approach to view problems as
parts of an overall system, rather than reacting to a specific
part, outcome, or event. It is about understanding and seeing
through the complexity and managing interdependency of
sectors, disciplines, and individual contributions. The process
encourages understanding of the big picture by sensing its
large-scale patterns and responding to them. We might benefit
by carefully crafting a process for defining problems and only
then designing collective solutions. Ultimately, this reflects
design thinking in bringing more healthful ingredients,
foods, and products into the kitchen and/or marketplace that
consumers demand, by taking into account the influences of
the overall food system, food environment, and community. A
systems approach will require multicultural representation to
come around a specific initiative to prioritize, focus, and lever-
age the strengths of individuals throughout the various sectors
and diverse disciplines of study.
Partnerships to Achieve a Common Goal around
Healthier Grain-Based Foods
According to Rowe et al., PPPs can create a collaborative envi-
ronment that maximize cross-disciplinary expertise among
government, academic, and industry researchers to help solve
complex research problems. Leveraging the proficiencies and
competencies of a broad range of stakeholder experts could
maximize available expertise and collectively achieve the best
research outcomes while extending and enhancing limited
financial and other resources. Figure 1 addresses 12 proposed
principles of PPPs, offered in no strict chronological order
because each partnership has unique characteristics.
It Starts by Defining Shared Value and Vision
Our vision is to bring together the various sectors, including
government, industry, nonprofit, academia, and activist
groups, to begin articulating the needs, desires, and demands
for targeting research dollars. The acquisition of research dol-
lars will help fill a void, which increases our ability as a society
to cost-effectively deliver healthier grain-based foods into the
marketplace. This is a vision of operation whereby sharing of
intimate information, goals, and objectives makes it challeng-
ing to establish a common shared value leading to a common
purpose, which drives this network of individuals toward a
common outcome and benefit for business and society.
Vision
Increase the availability of grain-based foods that are PHAD
with the intention of making the healthy choice the easy and
natural choice for citizens.

Figure 1 PPPs should do the following:
1. Have a clearly defined and achievable goal to improve the health of
the public
2. Articulate a clear statement of work, rules, and partner roles,
responsibilities, and accountability, to build in trust, transparency,
and mutual respect as core operating principles acknowledging
there may be deal breakers precluding the formation of an effective
partnership in the first place
3. Ensure that objectives will meet stakeholder partners needs, with a
clearly defined baseline to monitor progress and measure success
4. Considering the importance of balance, ensure that all members
possess appropriate levels of bargaining power
5. Minimize conflict of interest by recruiting a sufficient number of
partners to mitigate influence by any single member and to broaden
private sector perspectives and expertise
6. Engage partners who agree upon specific and fundable research
question(s) to be addressed by the partnership
7. Enlist partners who are committed to the long-term goals as well as
to the sharing of funding and research data
8. Along with government and the private sector, include academics
and other members of civil society as partners
9. Select objective scientific measurements capable of providing
common ground for both public and private sector research goals
10. Adopt research questions and methodologies established by
partners with no vested financial interest in them, ideally in the
precompetitive space
11. Be flexible and ensure ongoing transparent communications
12. Consider a third-party convener to ensure equality at the table,
clarify rules, establish operational guidelines, and specify funding
arrangements
Values
Values are integral in the roles that individuals play through
the group process and dynamics, clarity of objectives, mutual
trust, cooperation, and expected outcomes.
Why Is There a Need for a PPP?
Food and ingredient industries are uniquely positioned with
the technical expertise to make gradual product formulation
changes that are well accepted by consumers. These gradual
product formulation changes will not only improve the prod-
uct nutritional profile but also move consumers toward dietary
recommendations:
How can we fund research initiatives that guide food for-
mulation and product development to make it easier for
consumers to eat in a way that supports dietary guidance?
Is there a research approach that simultaneously encour-
ages new ingredient and innovative technology develop-
ment, supports a profitable industry business model, and
delivers products with enhanced nutritional value that
appeal to consumers?
What Are the First Steps?
New research is needed to support system-wide changes and to
assure continuity from science (theory) to consumers (prac-
tice) throughout the food delivery system. This integrated and
multifaceted process would deliver the following:
FOOD GRAINS AND THE CONSUMER | Grains and Health 65
Alignment among stakeholders through gauged,
coordinated, and prioritized research efforts that bridge
the gaps between research and consumer solutions
A sustainable infrastructure guided by industry profitability
while gradually improving the availability and consumer
acceptance of healthier food products
Targeted, prioritized research efforts that allow scientists to
address the food attribute gaps related to quality, cost,
convenience, and efficacy, translating science into food
Research that ranges from fundamental to product applica-
tion of functional foods and health, including enhanced
delivery of bioactives
Research that focuses on the successful development,
delivery, and enhanced consumer intake of healthier food
ingredients (this knowledge can provide insight on what
innovation opportunities are feasible)
Who Will Guide This Process?
Assemble the right minds
The PPP will be composed of an expanded network of
working partners across all sectors and disciplines of the
food supply, closely linked to consumer groups.
We are asking stakeholders to be a part of a PPP, which
will be formed to identify and to convene these partici-
pants and to solidify the project scope and the overall
approach.
Why Participate?
PPP members will be uniquely positioned to
steer the fundamental research path from the ground floor,
before it becomes available to the greater grain-based food
industry/community;
view firsthand the output of this fundamental and transla-
tional research, which can feed research and development
efforts to reach targeted customer segments such as schools,
restaurants, home, and retail;
leverage the resulting innovation and new processing
technologies.
How Will This PPP Function?
1. Key representatives, from across the food supply, will con-
vene to identify and prioritize research issues and gaps.
2. Meaningful collaboration will occur through application of
collective knowledge and targeted research.
3. Mutually beneficial collective solutions for business and for
society will result, ultimately positively influencing public
health outcomes.
What Makes This PPP Approach Unique?
This will be an innovative and a gradual shift in the way we
think about our food supply resulting in a shared and coordi-
nated approach to research with the following governing
guidelines:
66 FOOD GRAINS AND THE CONSUMER | Grains and Health
Open and safe environment to share ideas
Time to identify research gaps that limit availability of
healthier foods in the marketplace
Supporting infrastructure to implement the changes
Freedom to innovate and to conduct the targeted research
necessary to transform science into food
What Will Success Look Like?
Short term to midterm
Coordinated and comprehensive plan that identifies research
gaps through an integrated and gradual approach so that basic,
applied, and translational research can be targeted toward the
development and delivery of healthier grain-based foods that
deliver the attributes of a food product that are appealing to
consumers.
Long term
Sustainable infrastructure in place that supports a coalition
across the food supply chain, developing a unified strategy
that leads to increased availability of grain-based foods that
meet dietary guidance and resulting in a model that meets both
business and societal needs.
Overview of Foods Research and Public Health
Figure 2 summarizes the formation of partnerships/-
collaborative processes of convening the sectors and disci-
plines in the top portion of the diagram. On the far right
side, key consumer issues are identified and prioritized related
to food environments, such as schools, restaurants, or retail.
Moving to the left, specific food attributes for grain-based food
categories are then discussed relative to overcoming supply
chain barriers and challenges through product development.
The food attributes of grain-based foods should ideally meet
standards for PHAD by customers. Based on right to left move-
ment from the food environment, to food attributes, to supply
chain challenges, the research input is identified and targeted
through the supply chain and focused on key audiences in
schools, restaurants, and retail.
Based on the 2012 Whole Grain Summit, Minneapolis,
MN, five recommendations were established. Summit partici-
pants issued a call for a unified direction for the global grains
and health communities to
promote communication and unify leadership among
organizations in both grains and health,
encourage the delivery of consistent WG messages to
consumers,
support development of new products through targeted
research on both the health-related efficacy of WG and
technology for improved product development,
improve the consumer environment for choosing WG
foods through new product development by grain-based
food companies as a main means of increasing WG
consumption,
encourage wider adoption of governmental policies sup-
portive of an effective approach toward greater WG avail-
ability in the marketplace.
Follow-up from the meeting will be posted on the Grains for
Health Foundations website (Whole Grains Summit 2012
Events Page: http://grainsforhealth.org/news-and-events/
events/2012-whole-grain-summit).
Further discussions with representatives from industry, gov-
ernment, grain organizations, and academic institutions that
proposed research initiatives/projects include
I. standardized approaches and methods to WG research,
II. WG impact on the microbiome project,
III. community WG farm-to-fork intervention in New Ulm, MN.
Standard Protocol for WG Intervention Studies
of Intermediate Metabolic End Points (Proposed
Project I)
Purpose
The main objective is to develop a standardized protocol
whereby future intervention studies of WG and surrogate end
points would contain very specific research methods. Identify-
ing the appropriate methods of WG intervention studies would
allow for easier comparisons across studies of similar WG and
of studies that investigate different types of WG. Conducting a
meta-analysis would then be more easily performed to ade-
quately address the role of WG in prevention of chronic
disease.
What Problem(s) Does the Proposed Project Address
and for Whom?
The current body of evidence supporting the role that WG
plays in promoting health is largely composed of observational
studies. Intervention studies have yielded inconsistent results
when examining WG and health end points, such as insulin
resistance and Body Mass Index (BMI). The existing interven-
tion studies are not truly amenable to meta-analyses to help
address the issue of small sample size because of the variability
in the populations studied, variability in the WG interventions
(sources and quantity), and lack of standardization in measur-
ing surrogate endpoints. The end result is a lack of compara-
bility across many of the existing randomized trials.
Developing a standard protocol for WG intervention stud-
ies of intermediate metabolic end points would have a huge
impact on the scientific community. Researchers would be able
to pool individual study results together for stronger meta-
analysis. This would not only reduce the need for a single,
large-scale intervention but also save researchers time and
money.
Altogether, a better understanding of the role that WG plays
in health would ultimately benefit citizens and communities,
providing clearer nutrition education and implementation.
Why a PPP for This Project?
1. Does the research have the potential for significant public health
impact?
A clearer investigation of the role WG plays in health
could provide industry better direction in production,
delivery, and labeling of WG foods. This ultimately
 FOOD GRAINS AND THE CONSUMER | Grains and Health 67

OVERVIEW FOR

FOOD RESEARCH AND PUBLIC HEALTH

ERNM

GOV

EN
T
PR
N- O

NO

FIT
PARTNERSHIPS

ME WGS
DE

ACA RY

DUST
IN

SCIENTIFIC FOOD
FOOD
INPUT SUPPLY CHAIN ENVIRONMENT
ATTRIBUTES

GROWIN G

CONVENIENCE GAP PRACTICAL SCHOOLS

P R O D U C TI O N

EFFICACY GAP HEALTHFUL RESTAURANTS

MANUFACTURING

COST GAP AFFORDABLE HOME

CO NSUMER

QUALITY GAP DESIRABLE RETAIL

CONSUMER BENEFIT

HEALTHY CHOICE IS THE EASY CHOICE.

Figure 2 An overview of public and private partnerships for food research and public health.

provides the consumer with more informed and healthier

WG. Some grain groups focus on communicating the ben-
options. When the science gets better, everyone wins.
efits of enriched grains, some on the fiber benefits, and
2. Does it have implications across many different food and/or
some on WG. Building a stronger science base will allow
beverage categories?
for clarity in messaging to product developers, health pro-
Over the last decade, WGs have made their way in to
fessionals, and the consumer.
almost every food and beverage category (cereal, bread,
3. Is it precompetitive in nature and what does this mean for the
pasta, smoothies, grab and go salads, snack bars, crackers,
food industry?
etc.), and this is a trend that is forecasted to continue as
restaurants begin to incorporate WGs onto their menus. Yes, the question is precompetitive. The investigation
Additionally, the grains research community currently and subsequent inquiry would benefit any/all players in
struggles to deliver a consistent message on the benefits of the grain food industry. A stronger science base for the
68 FOOD GRAINS AND THE CONSUMER | Grains and Health
food industry would provide clarity in product develop-
ment, ingredient procurement, packaging messages/mar-
keting claims, etc. A common protocol for WG research
means less money spent on research that does not deliver
the final clarity that industry is looking for and more
mileage out of the money that is spent on individual
interventions.
Who
As suggested by Rowe et al. Partners with complementary
capability, capacity, resource access, and experience should
bear the risk appropriate to their contribution and share fairly
in the benefits from the research based on the value they bring
to the partnership. Partners might include the US Department
of Agriculture Agricultural Research Service (USDA-ARS) and
industry interested in clarifying the role of WG in health, the
Centers for Disease Control and Prevention (CDC) and/or the
National Institutes of Health, and leading academic research
centers, such as Tufts University and University of Minnesota.
Where
USDA-ARS research centers and National Institute of Food and
Agriculture external funding sources may help fund and par-
ticipate in this effort.
WG Impact on the Microbiome: PPP (Proposed
Project II)
Purpose
There is increasing evidence that microbial communities that
reside in the gastrointestinal tract play a crucial role in health
and disease. It is also becoming increasingly evident that diet,
among other factors, may be an important variable in the
establishment and maintenance of a microbial ecosystem that
supports and promotes health.
What are the characteristics of
a healthy microbiota?
PROJECT SCOPE
DIET
Genetics
Lifestyle
Life stage
Other
How does a whole grain diet
influence the gut microbiota?
Figure 3 The possible scope of a collaborative research project to assess the impact of whole grain foods on the gut microbial communities.
Why
A number of epidemiological studies have established that WG
consumption is associated with positive health benefits.
Although reductionist experimental approaches have provided
some insights into the components of WG that might explain
these associations, a thorough understanding of the impact on
WG consumption on host metabolism would be incomplete
without addressing the role WG plays in modulating host
metabolism through its impact on gut microbial communities.
This project has the potential to further our understanding
of the positive benefits of WG consumption to human health.
The project would have applications across several food cate-
gories (cereals, snacks, etc.). This research may strengthen the
evidence-based justification for increasing our consumption of
WG foods in the face of rising popular sentiments against
grain-based foods.
Who
The participation of both academia and industry will be essen-
tial to explore the benefits of WG on gut microbial
communities and their impact on human health. The food
industrys knowledge and experience in creating food products
with meaningful quantities of WG without compromising taste
would be an essential complement to the academic
institutions capabilities for assessing the impact of such
foods on gut microbial communities and health (Figure 3).
Community WG Farm-to-Fork Intervention in New
Ulm, MN (Proposed Project III)
Purpose
To create a healthful, sustainable local grain food supply
within a Minnesota community using a PPP model to leverage
diverse expertise to address WG intake. This proposal will focus
on WG foods in away from home (AFH) eating establishments.
Ecological and community-based participatory research
Gut
Microbiome HEALTH
How does the gut microbiota
influence health?

(CBPR) approaches will be utilized to increase WG availability,
sustainability, and consumption of AFH foods by increasing
the number of WG offerings and percent WG content in grain-
based foods found in restaurants and grocery store eateries.
This supply chain intervention will start in New Ulm,
Minnesota, as part of GrainUP!, a grassroots WG consumer
awareness campaign that encompasses the crucial industry-
facing supply chain work to create a road map for increasing
WG menu offerings AFH.
The proposed study will test the following hypothesis:
increasing the WG food offerings by 2550% (depending
upon the AFH setting and food category) and gradually intro-
ducing an increase in the WG content of popular grain-based
items in New Ulms AFH food environment (restaurants and
grocery store eateries). The hypothesis is that this will increase
AFH WG consumption from an average of one-third serving
per day to 1 serving per day, thus contributing to the Dietary
Guidelines for Americans overall recommended three daily
servings of WG. We aim to address availability, consumption,
and sustainability at a systems level.
Why
For almost a decade, the Dietary Guidelines for Americans have
recommended that Americans consume at least three ounce
equivalents (servings) of WG foods daily for optimal health.
WG foods improve heart health and weight management and
lower risk of certain cancers. Yet in spite of health benefits and
Dietary Guidelines for Americans recommendations, the average
American consumes only 0.8 servings per day. Low WG con-
sumption may be linked to the food environment and consump-
tion of foods outside of the home. Today, the average American
spends 42% of his or her food budget on AFH foods, which
constitutes about 35% of daily calories. These foods tend to be
lower in nutritional quality often high in calories, fat, sodium,
and cholesterol and offer less than one-third ounce of WG per
day. Eating AFH as little as one meal per week can lead to
approximately two extra pounds of weight each year for adults.
Modifying the food environment where adults frequently eat is a
promising approach to bringing about sustainable improve-
ments in nutrition behaviors at a population level so that when
consumers are faced with making food choices, the healthier
choice WG foods is the easier and preferred choice.
Recent research examining the relative healthfulness of
restaurant menus has shown that of 34 restaurants assessed
in New Ulm, fewer than 26% offered a WG bread option.
According to Rebecca Lindberg, researcher of Hearts Beat
Back: The Heart of New Ulm Project, feedback from restaurant
owners and chefs involved in trying to improve the healthful-
ness of their menu offerings has indicated that the WG category
is one of the most difficult menu items to change (oral com-
munication, 17 March 2014). Recent restaurant and grocery
store eatery assessments demonstrated that increasing AFH WG
menu choices remains difficult to influence when compared to
other healthful restaurant practices.
Why a PPP for This Project
1. Does the research have the potential for significant public health
impact?
FOOD GRAINS AND THE CONSUMER | Grains and Health 69
Yes, this has potential for significant public health impact.
Investigating the current barriers to a sustainable WG supply
chain consistently delivering healthier WG options to AFH
settings benefits the consumer. By allowing chefs, manu-
facturers, researchers, and policy makers to understand the
barriers, challenges, and opportunities to increasing WG
consumption AFH, steps can be taken to make the WG choice
the easy choice. This supports the Dietary Guidelines for
Americans recommendation of eating at least three servings
of WG a day.
2. Does it have implications across many different food and/or
beverage categories?
Grains are present in most AFH food categories, from
appetizers to dessert. In addition, their appearance usually
includes other food groups, such as fruits and vegetables,
meats and legumes, and dairy. Grains can serve as the
perfect vehicle for healthier foods AFH and provide an
opportunity to address portion sizes and added sugars
and fats.
3. Is it precompetitive in nature and what does this mean for the
food industry?
This proposal provides the opportunity for both ingre-
dient and consumer packaged goods manufacturers to
develop and pilot products and supply chain strategies, in
a precompetitive, collaborative environment. This applied
research model holds the potential to streamline processes,
reduce costs, and ensure technology transfer that can be
applied to other cities and communities.
Who
This proposal involves the resources and expertise of the Hearts
Beat Back Project (Minneapolis Heart Institute Foundation,
Allina Health, and the community of New Ulm, MN), food
industry, academics at the University of Minnesota, and project
management from the Grains for Health Foundation. The
Hearts Beat Back Project has previously received a USDA
Farmers Market Promotion Program grant, so has relationships
with farmers/producers, and is currently finishing a CDC Com-
munity Transformation grant (expanding their work to the rest
of Brown County, MN).
Where
The intervention will support current work of the Hearts Beat
Back Project, a 10-year population-based demonstration pro-
ject designed to deliver interventions at individual, institu-
tional, and community levels that reduce the rate of heart
disease risk factors in adult residents of New Ulm. Minneapolis
restaurants will be the next intervention site(s), once a sustain-
able supply chain approach and model is established, with
eventual rollout to other nationwide cities and communities.
When
Project work will occur over a 2-year period starting July 2014,
in the four phases listed in the succeeding text:
70 FOOD GRAINS AND THE CONSUMER | Grains and Health
Phase 1: Environmental scan for assessments and establishment
of AFH community forums
Phase 2: Gradual implementation of intervention using the
CBPR approach
Phase 3: Social marketing and promotion
Phase 4: Final assessments and evaluation of likelihood
intervention outcomes will be sustained
How
Typically, the last mile of the AFH food supply chain is where
many of the complications with WG foods reaching the con-
sumer occur. Given this, intervention will occur at various
points along the food supply chain (miller, baker, food
manufacturer, and AFH eating environment) to address WG
food quality, affordability, and desirability. An ecological
framework and the CBPR approach will be used to assess,
implement, promote, and measure a gradual increase in WG
offerings and percent WG content of current grain offerings.
WG foods of varying refined grain/WG content will gradually
be introduced in these establishments, along with technologies
to help reduce rancidity and enable tracking of shelf life of WG
flours to ensure the delivery of a fresh, safe, nutritious, and
palatable final product. The goal is that the increased consumer
acceptance and consumption of WG foods in AFH setting will
become permanent and sustainable.
Conclusions
New research is needed to support system-wide changes and to
assure continuity from science (theory) to consumers (prac-
tice) throughout the food delivery system. Public and private
research is one approach to explore ways of identifying
research initiatives that might help fill gaps in the food supply
chain, food environment, and community settings to more
successfully deliver healthier grain-based foods into the mar-
ketplace that consumers demand and readily consume.
A process is necessary to examine the issues, problems, and
gaps that exist relative to the development, delivery, and con-
sumer intake of healthier grain-based foods. Each research
project has merit as it relates to enhancing the development,
delivery, and consumption of grain-based foods made with
WG ingredients. Each project is complex in its own way and
affects different aspects of the food system and food environ-
ment along with community input and influence. We began to
address these research initiatives during a PPP meeting held on
34 December 2014, in Washington, DC. The outcomes of this
meeting will be discussed in another communication venue.
Exercises for Revision
Why are public and private partnerships needed to support
grain-based research?
What kind of thinking looks at parts of an overall system?
How is this helpful in grain-based research?
What is the benefit of a PPP in terms of solving complex
research problems?
Public and private partnerships combine what type of sec-
tors and disciplines?
Exercises for Readers to Explore the Topic Further
Identify the challenges and opportunities (gaps) related to
the substitution of WG wheat flour (50% whole wheat
flour) in a restaurant pizza. Track the pizza (1) through
the supply chain from field to fork and (2) in the food
environment, such as in a restaurant, and explore the char-
acteristics of the community patrons who frequent this
local restaurant.
How might a flour milling or packaged foods company use
this information related to the gaps in the food supply,
food environment, and community patrons?
What type of training programs might colleges, universities,
and industry consider in training of students and young
professionals to understand more about the food system,
food environment, and surrounding community who pur-
chase and use their products?
Further Reading
Ackoff RL (1981) Creating the Corporate Future: Plan or Be Planned for. New York:
Wiley.
Andersson A, Tengblad S, Karlstrom B, Kamal-Eldin A, Landberg R, Basu S, et al.
(2007) Whole grain foods do not affect insulin sensitivity or markers of lipid
peroxidation and inflammation in healthy, moderately overweight subjects. Journal
of Nutrition 137: 14011407.
Beauman C, Cannon G, Elmadfa I, Glasauer P, Hoffmann I, Keller M, et al. (2005) The
principles, definition and dimensions of the new nutrition science. Public Health
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 Food Grains: Intolerance, Allergy and Diseases
Contents
Cereal Allergens
Nutrition: Beriberi, A Deficiency Related to Grains
Celiac Disease
The Gluten-Free Diet
Cereal Allergens
AS Tatham, Cardiff Metropolitan University, Cardiff, UK
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Wheat, maize, and rice are major contributors to human
diets.
The cereals contain proteins (albumins, globulins, and pro-
lamins) that are implicated in food allergy and intolerance.
Four major types of wheat allergy are described, associated
with proteins that are part of protein superfamilies.
These superfamilies account for the cross-reactivity observed
between the cereals and between botanical species.
Currently, there are no treatments of cereal allergy (apart
from treating the attendant symptoms).
Patterns of cereal allergy are changing as patterns of cereal
consumption change.
Learning Objective
To achieve understanding of the types of cereal allergy,
allergens, clinical manifestations, and treatment.
Introduction
The most common allergic foods associated with
immunoglobulin E (IgE)-mediated reactions on a worldwide
basis are fish, peanuts, soybeans, milk, eggs, crustaceans, tree
nuts, and wheat. These foods commonly account for over 90%
of food allergies. Food allergies are increasing in prevalence and are
currently estimated to affect  28% of the population. The rea-
sons for this increase remain unexplained, as do the mechanisms
by which food proteins become capable of inducing an allergic
response in an individual. The prevalence of cereal allergies is low,
a figure of 2% of the population often being quoted.
Wheat allergy is defined as an adverse (IgE-mediated)
reaction to wheat, as distinct from intolerance (celiac disease),
which is a cell-mediated inflammatory reaction in the intesti-
nal lining. Wheat allergy can be divided into wheat-dependent
exercise-induced anaphylaxis (WDEIA), bakers asthma
72 Encyclopedia of Food Grains, Second Edition
(occupational asthma) (both of which have well-characterized
allergens), classic food allergy (affecting the skin and gastroin-
testinal and respiratory tracts), and contact dermatitis (where
the allergens are less well defined).
Wheat, rice, and maize constitute the big three cereal
crops, with the cereals forming the staple of many diets. It is,
therefore, perhaps unsurprising that cereal allergies, arising
either through ingestion or through exposure to cereal flours
and dusts, are increasing.
Symptoms and Pathology
IgE-mediated food allergies can elicit life-threatening reactions.
Allergy refers to an adverse immune reaction to food proteins
that are normally tolerated by the immune system. IgE acts as a
mediating antibody in the development of inflammatory
responses. These reactions, occurring in the lungs, skin, or
digestive tract, include asthma, rhinitis (hay fever), oral aller-
gies, food and contact allergies, and anaphylaxis; the most
severe form of allergic response involves a whole-body reac-
tion, which in some cases can lead to death.
The development of an allergic response involves several
steps. The first is sensitization, where the allergen stimulates the
production of antigen-specific IgE antibodies that can bind to
mast cells. This does not induce an allergic response, but repeated
contact with the antigen on subsequent ingestion binds to the
IgE, inducing the mast cells to release chemicals, such as hista-
mine, that cause the various symptoms of food allergy (inflam-
mation and anaphylaxis). Some individuals are referred to as
atopic, where they produce relatively high amounts of IgE and
are prone to becoming sensitized and developing an allergy.
Cereal Proteins
The proteins found in cereals fall into a limited number of
protein superfamilies that are characterized on the basis of
http://dx.doi.org/10.1016/B978-0-12-394437-5.00085-1
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Cereal Allergens
their highly conserved sequence regions, structures, and func-
tions and account for the cross-reactivity of allergens that are
found within the cereals. The prolamins (defined as aqueous
alcohol-soluble seed storage proteins) of wheat, barley, and
rye, and to some extent oats, are related; those of maize, millet,
sorghum, and rice are in part related but differ significantly
from wheat, barley, rye, and oats. Other protein families
include the pathogenesis-related (PR) proteins, which are
involved in the defense systems of plants against a host of
different pathogens, and also albumins and globulins that
can be involved in storage functions and metabolic proteins.
Understanding these relationships can be critical in under-
standing cross-reactivity in cereal allergy and intolerance.
Allergic reactions to wheat can be divided into inhalational
allergy (bakers asthma), ingestion (WDEIA), and also wheat
allergy and contact dermatitis, caused by skin contact with
wheat proteins. In bakers asthma and WDEIA, the allergens
are well characterized, whereas the allergens involved in wheat
allergy and contact dermatitis appear to be related to a range of
proteins that vary between individuals.
Bakers Asthma
Bakers asthma is one of the most common forms of occupa-
tional asthma, caused by the inhalation of wheat and cereal
flours and of dusts and other substances commonly found in
bakeries and flour mills. In bakery and flour-handling workers,
studies indicate 510% suffer from asthma and 1020%
from rhinitis, making it one of the most common occupational
allergies.
Studies have shown that multiple allergens are present in
cereal flours that are recognized by IgE from patients sera. One
group, the a-amylase inhibitors, contains the major allergens,
with other allergens (including the lipid-transfer proteins
(LTP), peroxidase, thioredoxin, and serine proteinase inhibi-
tors) showing the strongest IgE reactivates in wheat flour; the
prolamins, thaumatin-like proteins, and acyl-CoA are also
implicated. In addition, a range of flour improvers/additives
contribute to the allergenic potential of cereal flours.
The a-amylase inhibitors consist of 1216 kDa polypep-
tides with 45 intrachain disulfide bonds, which are tightly
folded, making them thermally stable and difficult to digest.
They are encoded by multigene families in wheat, barley, and
rye, giving rise to marked heterogeneity in their composition.
In addition, they can occur as monomers, homodimers, and
tetrameric aggregates. They are active against the a-amylase of
insects and in the cereals are active in plant defense. Homology
between the proteins found in wheat, barley, and rye accounts
for the cross-reactivity between the flours from these cereals.
They are panallergens, being present in the flours of other
cereals.
In addition, LTPs constitute a family of panallergens; their
main function is in plant defense against phytopathogens
(fungi and bacteria); they are classified as PR proteins. They
have been identified as the major allergens in many fruits in
the Rosaceae family (e.g., peach, plums, and apples). In bakers
asthma, they are major allergens.
In addition to the water/salt-soluble allergens, which con-
stitute the major classes of allergens in bakers asthma, the
prolamins are also implicated. Patients sera have shown
73
binding to the a-, g-, and o-type gliadins and low-molecular-
weight subunits of glutenin.
Detailed studies of sequentially extracted wheat flour, into
salt-soluble (albumins and globulins) and alcohol-soluble gli-
adins and glutenins, indicate that patients sera recognize mul-
tiple allergens.
Other ingredients are used as flour/bakery additives, often
starch-cleaving enzymes, and some of these have been found to
be allergenic. Fungal a-amylase from Aspergillus oryzae is added
to digest starch and is allergenic, as are cellulases and xylanases
from Aspergillus niger, added to flour to digest fiber. Soy flour,
added to wheat flour as a source of lipoxygenase to whiten
flours, is responsible for food and inhalational allergies. Soy-
bean allergy has been implicated in bakers asthma.
Wheat-Dependent Exercise-Induced Anaphylaxis
Food-dependent exercise-induced anaphylaxis (FDEIA) is
defined as when food intake, followed by exercise, results in
the display of allergic symptoms (from urticaria to anaphylac-
tic shock). When food intake and exercise are taken indepen-
dently of each other, no symptoms are observed. Ingestion of
the causative food and exercise within 23 h (moderate to
strenuous) are required for the allergic response to occur.
Wheat is reported, along with crustaceans, to be one of the
foods most commonly reported to be associated with FDEIA.
The incidence of FDEIA is difficult to evaluate but estimated to
account for 515% of all anaphylactic cases; of this, one-third
to one-half are estimated to arise from FDEIA. In a Japanese
study, the incidence was found to be 0.017% and 0.031% for
cases of FDEIA and EIA, respectively.
The major allergens associated with WDEIA have been
identified as the o5-gliadins (from the B genome of wheat)
and the high-molecular-weight (HMW) subunits of wheat. In
one study,  80% of patients had IgE reacting to o5-gliadins,
the remainder to HMW subunits. The g-35 and g-70 secalins of
rye and the g-3 hordeins of barley cross-react with wheat
allergic sera, but no cross-reactivity with the avenins of oats
has been reported.
In wheat, the epitopes (amino acid sequences that cause the
allergic response) are well characterized. Four immuno-
dominant epitopes in the o5-gliadins of wheat have been
identified (QQIPQQQ, QQFPQQQ, QQSPEQQ, and
QQSPQQQ). Mutational analysis, using synthetic peptides,
indicated that Gln at positions 1, 5, 6, and 7 and Pro at
position 4 of the heptapeptides were critical for IgE binding.
Omega gliadins, derived from the A and D genomes of wheat,
do not contain these epitopes and are not implicated in
WDEIA. Three IgE-binding epitopes, in the repetitive domains
of the HMW subunits of wheat, have been identified, QQPGQ,
QQPGQGQQ, and QQSGQGQ, using synthetic peptides.
Aspirin increases the levels of gliadin-derived peptides cir-
culating in the bloodstream and upregulates the allergic
response to specific foods, including wheat. Wheatexercise
and wheataspirin challenges increased the levels of gliadin-
derived peptides in the blood, indicating that both exercise and
aspirin facilitate allergen absorption from the intestine.
The specificity of the association between o5-gliadin and
WDEIA has led to the development of diagnostic tests measur-
ing the levels of IgE directed against recombinant o5-gliadins.
74 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Cereal Allergens
Wheat Allergy
Wheat allergy caused by the ingestion of wheat proteins can
give rise to a range of symptoms including urticaria, atopic
dermatitis, and anaphylaxis. Wheat allergy in children, similar
to egg and milk allergies, is outgrown by the majority of
children by the age of 35. In children, the most common
symptoms are atopic dermatitis. In adults, wheat allergy is
less frequently observed.
Studies have shown that the IgE fractions for both adults
and children react with gliadin (a-, g-, and o-types) and glute-
nin fractions, albumins, and globulins, the latter two groups
being strongly correlated with atopic dermatitis. Similar stud-
ies have shown differences in the pattern of IgE recognition
related to age and symptoms. Patients who showed anaphy-
lactic reactions, similar to those with WDEIA, had IgE to the
o5-gliadins.
Patients with this form of allergy often show IgE to multiple
allergens, and diagnosis, other than by food challenge, is prob-
lematic. Wheat-specific IgE concentrations do not appear to be
good indicators of the outcomes of provocation (food chal-
lenge) tests, so even with a low or negative outcome, wheat
allergy cannot be excluded without a challenge test, if clinical
symptoms suggest wheat allergy.
Contact Dermatitis
Contact dermatitis is an inflammation of the skin caused by
coming into direct contact with an allergen. This form of wheat
allergy mainly occurs in bakery workers and others coming
into occupational contact with wheat flours and dusts. More
recently, contact dermatitis has been reported from cosmetics
where hydrolyzed wheat proteins have been used as an ingre-
dient to improve hydration properties, although their
occurrence is uncommon. Oats, also used as an ingredient in
cosmetics, have also been reported to cause contact dermatitis.
There are few studies on the allergens responsible; a-amylase
inhibitors, albumins, and globulins and gliadins have all been
implicated.
Rice Allergy
Rice allergy is common in Eastern Asia where rice forms a
staple in the diet; it is less common in western countries. The
most frequently observed symptoms are atopic dermatitis,
eczema, asthma, and less commonly anaphylaxis. There are
few reports of rice allergy, and little is known of the incidence
or prevalence of rice allergy in Asian countries. Considering the
importance of rice in the diet of a large part of the worlds
population, this allergy lacks detailed studies on prevalence.
The major allergens are the a-amylase inhibitors, which are
implicated in 90% of patients exhibiting food-related atopic
dermatitis. Rice-induced anaphylaxis has been linked to the
LTP proteins, and cross-reactivity has been reported with LTPs
from peach and apple. Rice is also linked to occupational
asthma and contact dermatitis in workers handling rice and
rice-derived products.
Maize Allergy
Maize allergy has been reported in countries where maize
forms a staple in the diet, such as Africa, Central and South
America, and southern Europe. In the United States, the prev-
alence of allergy to maize and maize-derived sweeteners is low
(estimated at 0.016% of the population); the incidence else-
where is thought to be higher. Maize allergy is linked to occu-
pational asthma and contact dermatitis through handling
maize flour and food allergy by maize ingestion.
The major allergens are LTPs and a-amylase inhibitors; the
LTPs show cross-reactivity with LTPs from barley and rice;
minor allergens include thioredoxins, g-zein (an alcohol-
soluble seed storage protein), and trypsin inhibitors.
Allergies to Minor Cereals and Pseudocereals
Food and inhalational allergies have been reported for millets
and sorghum and for the pseudocereals buckwheat, amaranth,
and quinoa. These crops are recommended for inclusion in
gluten-free diets for celiac sufferers.
Effects of Food Processing on Allergenicity
Cereals are often consumed after being subject to a variety of
processing conditions, which may alter the epitopes in the
proteins responsible for their allergenic properties. Processing
can result in a reduction, no change, or an increase in the
number of epitopes in the protein or an increase in their
stability to digestion.
The effects of thermal processing on wheat proteins have
been reported, with different conclusions. Some studies have
reported a reduction in allergenicity on heating extracted iso-
lated proteins in solution or extracting them from heated glu-
ten and analyzing their IgE binding. Others have reported
similar IgE reactions from wheat proteins extracted from flour
then heated and cooled or from flour and water heated and
cooled and then extracted. In food challenges, some patients
responded to a lower dose of cooked flour than unprocessed
flour, which suggested an increase in the number of allergen
epitopes.
The effects of proteolysis (enzymatic digestion) to produce
hypoallergenic wheat have been developed, using enzymes
that digest known epitopes; however, these seriously affect
the functional properties of the proteins. Severe allergic
responses have been shown in hydrolyzed wheat proteins
used in cosmetics, where the unmodified protein did not
induce the response. This indicates that the process may intro-
duce epitopes. In addition, deamidation of the glutamine res-
idues of the prolamins may increase their allergenic potential;
it is known that deamidation of glutamine residues in the
celiac-active motifs enhances their activity.
Allergic responses have been reported from beers brewed
from barley and wheat and from barley malt. The major aller-
gen was identified as an LTP, which retained allergenicity after
being unfolded and chemically modified.
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Cereal Allergens
Diagnosis of Cereal Allergy
If a cereal allergy is suspected, a number of tests can be per-
formed for an accurate diagnosis.
Skin prick tests (SPTs) involve applying allergens, in solu-
tion, subcutaneously. In an allergic individual, these will pro-
voke mast cells to release histamine and produce inflammation
within about 15 min. Blood tests measure IgE levels in vitro in
blood serum. These tests measure sensitization, and not nec-
essarily a clinically relevant allergy. The most reliable tests
involve food challenges, which are used for immediate wheat
allergy detection. Although diagnostic in vitro IgE and SPT
assays are known to be unsatisfactory, they are extensively
used. The CAP system has a predictive value for wheat of
< 75%. The recent use of recombinant o5-gliadin for the diag-
nosis of WDEIA patients was reported to be around 80%, and
children with high concentrations of IgE specific to o5-gliadin
had a higher risk of failing a wheat challenge. The tests may, in
some part, be unsatisfactory due to the allergen mixtures that
are used. The albumins and globulins are relatively easy to
extract from wheat flour; the prolamins are less so, due to
their poor solubility in aqueous media and possible presenta-
tion in the IgE-binding assay.
Safe Amounts of Wheat Proteins for Patients with
Wheat Allergy
There is an inherent difficulty in determining the minimum
eliciting doses that will cause an allergic reaction. Individuals
will differ in their sensitivity to different allergens, and often,
multiple allergens are present; a combination of these may be
required to elicit a response. In children with wheat allergy,
810 g flour has been shown to induce symptoms, whereas in
another study, it required < 2 g; children generally have a lower
tolerance than adults due to their smaller body mass. WDEIA
appears to be triggered by larger amounts of flour (60 g), which
may be related to the allergen being present in small amounts,
but there are few reported studies. There are ethical issues with
feeding cereal flours to individuals with wheat allergy (as you are
inducing a clinical response), but further study is required.
Management of Allergies to Cereals
Managing cereal allergy involves reducing the risk of an allergic
reaction by avoiding the allergen and by medical treatments to
reduce symptoms.
Allergen avoidance is an essential step in managing food
allergies, for which there are currently no cures. Allergen label-
ing on packaged foods is a legal obligation in many countries.
However, the use of may contain places the onus on the
customer to make a decision whether to purchase and con-
sume the food. There is an increasing market for free-from
food products, especially gluten-free, providing a wide range
of products with similar textural properties to wheat-
containing products. Hidden allergens remain a problem,
where wheat gluten is used in a wide range of products. Studies
75
suggest that wheat gluten and wheat starch are found in more
than 30% of packaged supermarket foods.
Allergy medications control the symptoms and reactions to
allergens but do not cure the allergy. Antihistamines are the
most common form of allergy medication. During an allergic
reaction, the immune system releases histamine, which starts
the cascade effect of allergy symptoms, including swelling or
edema, a drop in blood pressure, and itching. Antihistamines
work by blocking the action of histamine, and they work best if
taken before an allergic reaction has started, but they can block
the further release of histamine after a reaction has started.
Emollient lotions and creams, used for eczema and dry skin,
may contain antibiotics to reduce the risk of infection and
steroids to reduce inflammation. Adrenaline is used as an
emergency treatment for anaphylactic shock, where sudden
high levels of histamine released during the allergic reaction
cause difficulty in breathing. Adrenaline alleviates the symp-
toms of the reaction and can be delivered by an autoinjector
before further emergency treatment.
Application of Genetic Modification
The use of genetic engineering to downregulate gene expres-
sion has been demonstrated in a number of cereal crops,
particularly in relation to reducing the levels of specific pro-
teins that contain celiac disease-activating epitopes. These
approaches have the potential to reduce allergens in cereals.
Hypoallergenic rice has been developed with downregulation
of the gene family encoding a-amylase inhibitors, which are
major food allergens in Japan. These approaches reduced the
levels of the inhibitors down to about 20% of their levels in
control plants. Wheat has been transformed to silence the o5-
gliadins (active in WDEIA), resulting in complete or substan-
tially reduced levels relative to untransformed controls, with
the retention of technological properties. These approaches
could be used to reduce allergenicity by introducing stable
heritable changes in the cereal genome.
Conclusions
The major allergens associated with bakers asthma and
WDEIA have been identified. The a-amylase inhibitors are
most important in bakers asthma but less important in food
allergy. The prolamins appear important in food allergies,
particularly the o5-gliadins in WDEIA, and less so in bakers
asthma. Several other minor allergens have been identified and
their clinical relevance needs to be determined. There is a lack
of information relating to rice allergy and to a lesser extent
maize allergy. With the wider use of minor cereals and pseu-
docereals in the diets of cereal-allergic patients, it might be
expected that there will be an increase in the incidence of
allergy to these cereals as they become established foodstuffs.
More reliable diagnostic methods are being developed for
cereal allergies based on the major allergens, and advances are
being made for their removal/reduction in transgenic crops.
The wider availability of free-from and gluten-free will give
the allergic consumer greater choice and, with legislation
76 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Cereal Allergens
related to labeling, greater confidence in the foods that they
consume.
Exercises for Revision
What are the major types of allergy associated with wheat?
Why are patterns of cereal allergy changing worldwide?
What are the major classes of proteins in cereal grains?
What are the main types of wheat allergy and their causative
allergens?
Exercises for Readers to Explore the Topic Further
Which groups of proteins account for cross-reactivity
between botanical species?
What are the potential uses of GM technology in reducing
cereal (and general) food allergies?
What are pseudocereals? How can they contribute to the
diet of cereal allergy and intolerance sufferers?
Further Reading
Altenbach SB and Allen PV (2011) Transformation of the US bread wheat Butte 86 and
silencing of omega-5 gliadin genes. GM Crops 2: 6673.
Gilissen LJWJ, van der Meer IM, and Smulders MJM (2014) Reducing the incidence of
allergy and intolerance to cereals. Journal of Cereal Science 59: 337353.
Hischenhuber C, Crevel R, Jarry B, et al. (2006) Review article: Safe amounts of gluten
for patients with wheat allergy or coeliac disease. Alimentary Pharmacology and
Therapeutics 23: 559575.
Inomata N (2009) Wheat allergy. Current Opinion in Allergy and Clinical Immunology
9: 238243.
Matsuo H, Kohno K, Niihara H, and Morita E (2005) Specific IgE determination to
epitope peptides of omega-5 gliadin and high molecular weight glutenin subunit is
a useful tool for diagnosis of wheat-dependent exercise-induced anaphylaxis.
Journal of Immunology 175: 81168122.
Pastorello EA, Farioli L, Pravettoni V, et al. (2009) Maize food allergy: Lipid transfer
proteins, endochitinases, and alpha zein precursor are relevant maize allergens in
double-blind placebo-controlled maize-challenge-positive patients. Analytical and
Bioanalytical Chemistry 395: 93102.
Salcedo G, Quirce S, and Diaz-Perales A (2011) Wheat allergens associated with bakers
asthma. Journal of Investigational Allergology and Clinical Immunology 21: 8192.
Sheffer AL and Austen FA (1980) Exercise-induced anaphylaxis (1980). Journal of
Allergy and Clinical Immunology 66: 106111.
Tada Y, Nakase M, Adachi T, et al. (1996) Reduction of 1416 kDa allergenic proteins in
transgenic rice by antisense gene. FEBS Letters 391: 341345.
Tatham AS and Shewry PR (2008) Allergens in wheat and related cereals. Clinical and
Experimental Allergy 38: 17121726.
 Nutrition: Beriberi, A Deficiency Related to Grains
KJ Carpenter, University of California, Berkeley, CA, USA
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 318325, 2004, Elsevier Ltd.
Topic Highlights
Beriberi, a name meaning great weakness is due to B-vitamin
(thiamin) deficiency, commonly associated with milled rice
consumption.
White rice alone, absent a diverse diet providing varied
sources of vitamins, is severely deficient in thiamin.
There is no easy, convenient food source that provides high
thiamin. Parboiling of rice is beneficial, as is enriching rice
with synthetic vitamins.
Learning Objectives
To understand the etiology of beriberi and the history of its
dicovery.
To understand the effect of grain processing on thiamin
levels and hence on propensity to induce beriberi.
To understand the effects of other dietary habits, e.g. exces-
sive alcohol consumption, on thiamin deficiency.
Introduction
Grains have been the major source of both energy and protein
for the worlds population for millenia. However, it is now
realized that, when the diet lacks a variety of other foods, the
grains are also needed to supply a considerable number of
other essential micronutrients if those eating them are to
remain healthy. But the processing of grains to remove the
bran and germ, in addition to its favorable effects of increasing
shelf life by removing much of the oils that can become rancid,
and providing foods with a smoother feel by removing most of
the fiber, does also cause the loss of a large proportion of their
other nutrients.
It now appears that the emergence of the disease pellagra
amongst communities in the southern states of the USA from
around 1910 is explained by the introduction of machinery
that allowed the replacement of stone-ground whole corn
meal with de-germed meal, and that this, combined with
white wheaten flour and little in the way of lean meat, eggs,
and milk, resulted in a diet deficient in the vitamin niacin.
Until this came to be understood in the 1930s, several thou-
sand people died and even more suffered a miserable and
humiliating disease.
The following article summarizes the way in which another
disease, beriberi, that was causing huge suffering in the Far
East, particularly in the earlier period 18701910, came to be
understood as the consequence in most cases of people having
polished white rice as their staple food with little in the way of
supplementary foods. It was the work of the pioneers
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00080-2
struggling to understand this disease that led to the discovery
of the class of micronutrients that now is known as vitamins.
It is a complicated story with many false leads, but it pro-
vides a valuable lesson in how science does eventually advance,
and the dangers of over-simplification of the problems of the
real world. It also shows how it finally proved possible, as a
result of collaboration between industry and academia, to
enrich highly milled grain products so that they still pro-
vided a good contribution of vitamins and minerals.
Occurrence in Asia
The first Western physicians allowed to work in Japan in the
1870s were surprised to discover the existence of a serious
disease previously unknown to them and second only to
smallpox in its ravages. In Japan, it was known as kakke,
but was soon recognized as being identical to the disease
known in South-east Asia as beriberi, a native name now
universally adopted, which may originally have meant great
weakness. Characteristically, it began with a feeling of weak-
ness in the legs and a loss of feeling in the feet. Then, in many
but not all cases, the legs and then the trunk would swell with
retained water. Finally, the heart would be affected so that the
subject gasped for breath, and would die from heart failure.
Older records from both Japan and China showed that it
had been known for some centuries, although it had been the
opinion of two eighteenth century Japanese physicians that the
disease had become worse after
1750. The early records also
indicated that it was largely a disease of the wet summer
months and could attack even the well-off.
Infection or Malnutrition?
Those most at risk were men in the newly modernized Japanese
army and navy, and also prisoners. As these were all people
living together in large groups, and with the excitement in this
period at other diseases being traced to the transmission of
pathogenic bacteria, this seemed a likely cause for beriberi also.
Yet, it was difficult, on this basis, to explain the frequent
observation that a naval ship would leave its base with all its
crew in good health, yet, after a month or more in isolation at
sea, the disease would sweep through the crew.
Kanehiro Takaki, a surgeon on the naval staff, was directed
in 1878 to work on the problem. He knew that the ships had
been built in Britain and that they followed the general prac-
tices of the British navy where there was no beriberi. The only
difference that caught his attention was in the rations issued to
the men: the Japanese issues contained less protein and did not
meet the high standard in force at that time in Europe. He
therefore persuaded his superiors to permit a trial of modified
rations with a proportion of the rice being replaced by meat,
77
78 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Nutrition: Beriberi, A Deficiency Related to Grains
condensed milk, vegetables, and barley. The change was a
complete success, and it was found that even just the use of
barley in place of one half of the rice staple was enough to
prevent the disease, which Takaki now believed to have
resulted from a deficiency of protein in the earlier rations.
The Japanese army, perhaps as the result of interservice
rivalry, did not follow the navy and in the short Russo-Japanese
war of 190405, some 100 000 of their soldiers had to be
invalided home from Manchuria suffering from beriberi.
A Disease in Chickens
Meanwhile, the disease had become an equally serious prob-
lem in the Dutch East Indies (now Indonesia) (Figure 1). After
a punitive military expedition had to be withdrawn because of
a beriberi epidemic, the Dutch government dispatched a small
commission to try to identify the bacteria responsible for the
disease. After a few months, it was thought that the microor-
ganism had been found, and Christiaan Eijkman, a young
Army physician, remained behind to confirm its activity in
animal models.
Some of the chickens that Eijkman had injected with blood
from beriberi patients developed signs of leg weakness, but so
did some of his uninjected controls, suggesting that the condi-
tion was so infectious that it could jump from cage to cage.
Autopsies of the affected birds showed degenerated peripheral
nerves. But in the following months, none of the next batch of
birds developed the condition. Eijkman discovered that when
the leg weakness had appeared, the man in charge of the birds
had been feeding them on cooked white rice left over from
feeding the beriberi victims in the adjoining hospital, instead
of buying rough, feed-grade rice.
Figure 1 Two prisoners in Java with beriberi and needing assistance
to walk. (Reproduced with permission from Vorderman (1897)
Onderzoek naar het gevangenissen op Java en Madoera en het
voorkomer van beri-beri order de geintemeerden. Batavia: Jav. Boekh &
Drukkerij.)
A long series of feeding trials confirmed that birds fed on
white rice would become sick with leg weakness, whereas those
given supplements of rice polishings (still present in feed-grade
rice) remained healthy. This was only an animal disease, but a
survey by the medical inspector of prisons in Java showed that
prisoners who had been receiving white rice as their staple
issue were susceptible to beriberi, whereas those receiving
brown rice were not.
The Concept of a Vitamin
Eijkman, who believed that the disease was a kind of starch
poisoning, now had to be invalided home with malaria. His
successor, Gerrit Grijns, found that birds became sick even
when fed on meat that had been autoclaved. After further
work, his statement in 1901 was perhaps the progenitor of
the vitamin era in nutritional research: there occur in vari-
ous natural foods substances which cannot be absent without
serious injury. . .they are easily disintegrated. . .and cannot be
replaced by simple chemical compounds.
The work of Eijkman and Grijns was confirmed by British
investigators in Malaysia and by Americans in the Philippines,
and attempts began to extract the active material from rice
polishings and to concentrate it. There are moving accounts
of scientists in Manila being implored by local doctors to bring
a few spoonfuls of extracted syrup to save the lives of infants
with beriberi, and of the babies spectacular recoveries. Women
themselves seemed less susceptible to beriberi than men, but
when mothers were receiving a diet of low thiamin content,
their breast-fed babies were at a high risk of dying with acute
infantile beriberi.
Isolation of Thiamin
Isolation of the active factor proved very difficult. Each stage
of extraction, and then further partitioning by reprecipita-
tion involved biological assays with birds. Eventually, the
next generation of Dutch workers in Indonesia was success-
ful, obtaining a few milligrams of active crystalline material
after starting with one-third of a tonne of rice polishings
and going through at least 16 separation stages in which
much of the vitamin was lost. It was found that adding just
2 ppm of the crystals to white rice was enough to keep birds
healthy. In 1931, the crystals were found to contain sulfur as
well as carbon, hydrogen, nitrogen, and oxygen, and the
chloride salt was shown to have the empirical formula
C12H18N4SO2Cl2.
There were, of course, almost innumerable ways in which
these atoms could be combined. By good fortune, Robert R.
Williams in the USA found that adding sodium sulfite to a
solution of the vitamin led to its division into two roughly
equal halves. Further work at a number of centers showed that
one of these compounds contained a pyrimidine and the other
a sulfathiazole ring. By 1937, a synthesis of the active molecule
was achieved. It was named thiamin or thiamine (i.e., the
sulfur-containing vitamin) and soon began to be produced and
marketed as a pharmaceutical.
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Nutrition: Beriberi, A Deficiency Related to Grains 79

The Analysis of Foods
Thiamin can be oxidized to a highly fluorescent derivative,
thiochrome. This property is used to measure the thiamin
contents of different foods, even at levels of less than 1 ppm.
The procedure is specific, and no other naturally occurring
compounds have been found that give the thiochrome reaction.
However, thiamin can react with polyphenols and a compound
present in garlic to give derivatives that are still biologically
active (as will be referred to again) but do not give the thio-
chrome reaction. The analytical procedure may therefore under-
estimate the efficacy of a product. It is essential, therefore, to
have a confirmatory bioassay before knowing for certain that
any kind of processing has caused significant loss of thiamin.
Rice and Other Staples
Figure 2 illustrates the thiamin levels in the worlds major
staple foods, both when fully milled and when minimally
processed. In the case of grains, the latter means removal of
the husk (or hull) but no more. It is clear that, for each grain, the
full milling that removes both the bran and germ results in the
loss of a major portion of the thiamin originally present.
White rice is not that much lower than white wheat flour in
its content of the vitamin, but after the grains have been
prepared for consumption, the difference is increased. White
rice is normally washed several times, and this alone can
remove half the thiamin present, and boiling in excess water
can again halve the level of remaining vitamin. In contrast,
white wheat flour is most commonly baked into bread with
yeast as the raising agent, and this causes little loss of thiamin.
There is no evidence that cooked white rice has any posi-
tively harmful qualities, but if it is the major item in a diet that
contains only small amounts of foods that are richer in thia-
min, so that the diet as a whole provides no more than about
0.25 mg per 1000 kcal, it is not surprising that beriberi should
gradually develop.
The data in Figure 2 also explain the Japanese experience
that serious problems with beriberi in their navy in the late
1800s disappeared when one-half of their rice ration was
replaced by barley.
The same figure also shows the low thiamin content of
tapioca prepared from cassava roots. This explains the exis-
tence of beriberi in Brazil at the same period among even
well-off people whose favorite foods were tapioca and molas-
ses. Their preferred protein supplement was dried, salted cod,
which had to be soaked for several days to leach out most of
the salt, which also removed most of the vitamin.
The very first reports of beriberi to reach Europe came from
Portuguese priests working in the Molucca Islands (at the
Eastern end of Indonesia) in the 1500s. Their staple was the
locally produced sago meal, now realized to be almost pure
starch, and the priests correctly attributed their weakness to a
lack of something in this food and asked to be provided by
their superiors with wheat flour.
Beriberi was also a serious problem in early spring in iso-
lated communities in Newfoundland in the early years of the
twentieth century. Their families, who would be cut off for
the winter, had to buy 6 months of supplies, with white flour
as their staple. They were also apparently not familiar with
using yeast to leaven bread, but cooked their flour with baking
soda, and it is known that much of the thiamin present
would be destroyed under the alkaline conditions during this
procedure.

Figure 2 Representative analytical values for the thiamin content of different staple foods: (1) after husking only ( ) and (2) after full processing
( ), as explained in the text, and also for some staple root crops, etc. Sago meal is not shown, as it contains only an insignificant level of thiamin.
(Reproduced with permission from Carpenter KJ (2000) Beriberi, White Rice and Vitamin B. Berkeley, CA: University of California Press.)
80 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Nutrition: Beriberi, A Deficiency Related to Grains
The Improvement of Rice
Once the association of beriberi with white rice in Asia had
been established, attempts were made to replace it in some way
with other foods. As already mentioned, barley was an eco-
nomic and well-accepted alternative in the diet of the Japanese
armed forces.
In the Philippines, a proposal was made to enforce the use
of brown rice by the local military. This, of course, is rice from
which the husk has been removed but not the entire bran layer
and germ (embryo and scutellum) (Figure 3). This was the
traditional staple of villagers in South-east Asia who had no
access to mechanical rice mills. They would pound their paddy
(i.e., rice still in the husk) in some kind of bowl and then
winnow the product so that the lighter husks blew away, and
the grains fell in a pile.
This procedure was time-consuming but created no prob-
lems when only enough was pounded for immediate use in the
next 24 h. However, it was repeatedly found that in the tropics,
brown rice on storage would become infested with insects of
different kinds, and the oil in the bruised germ would become
rancid. Since large organizations, or an army on the move,
needed large-scale supplies ready for cooking, brown rice did
not provide a practicable staple.
The early workers who discovered the association of beri-
beri with white rice had assumed that the important micronu-
trient was concentrated in the bran of the grain. However, it
was later realized that more was present in the germ area
(Table 1). Japanese millers then attempted to modify their
machinery so as to remove the bran without removing the
germ from the grain. The so-called germ rice that they were
able to produce proved to be both palatable and an improved
source of thiamin. However, millers were only able to produce
Figure 3 Dissection of a dehusked rice grain. (Reproduced with
permission from Carpenter KJ (2000) Beriberi, White Rice and Vitamin B.
Berkeley, CA: University of California Press.)
it with certain varieties of rice, and not with the bulk of the rice
favored in Japan.
A traditional method of processing rice common in Bengal
is called parboiling. It had been found that if rice in the husk
were to be steeped for a period in hot water and then allowed
to dry in the sun, the husks cracked off more easily on pound-
ing, and there was less breakage of the grains. Broken grains
had a lower commercial value.
In Malaysia, where many immigrant groups were employed
as laborers but living on their habitual diets, it was realized in

1910 that Bengalis were remarkably free from beriberi, and
studies in a mental hospital showed that replacing ordinary
white rice with polished rice prepared from parboiled grains
relieved the inmates from the disease. After thiamin had
been identified, analyses showed that the initial soaking of
the grain caused thiamin to diffuse into the endosperm and
to remain there as the grains dried out.
Again, this does not appear to be something that could be
applied more widely. The traditional soaking and sun-drying
leaves the rice with a characteristic musty flavor and slight
discoloration, which is acceptable only to people who have
grown up with it. The process can be modernized, with con-
trolled autoclaving and vacuum-drying for so-called conver-
sion of the rice, but this makes it too expensive for mass
consumption in developing countries.
The final procedure for the production of enriched rice is
to fortify it with the synthetic vitamin. For a flour, such as white
wheat flour, this is relatively simple, requiring only very careful
mixing of the traces of vitamin with a small quantity of flour,
then the blending of the premix to larger batches. This is
familiar, and indeed compulsory practice, in the USA and
UK, together with other vitamins and trace minerals. As a
powder, thiamin cannot be blended with grains of white rice.
However, methods have been developed of preparing vitamin-
rich granules with the size and appearance of rice grains, and
blending these in the ratio of one granule to 200 grains, so that
the mix has at least the thiamin content of brown rice. To
reduce loss of the vitamin during washing and cooking, the
granules are coated with a nontoxic resin at the final stage of
their production. This method of enrichment has been tested
in an area of the Philippines where beriberi was endemic and
has proved successful in greatly reducing its incidence. Unfor-
tunately, where widely separated villages each had a small
electric mill, there were practical problems in persuading
millers to pay for the premix when the product appeared
unchanged, but the price had to be a little higher.
Supplementing Foods
Unfortunately, there is no convenient food that is extremely
rich in thiamin. Dried brewers yeast contains 15 mg per 100 g,
but many people cannot tolerate it in more than extremely
small regular doses. Of the meats, pork is richest with lean
pork containing 1 mg per 100 g. Beef has only about one-tenth
as much. Dry peas and beans contain about 0.5 mg per 100 g.
Potatoes are another useful supplement on a dry matter basis,
they have slightly more thiamin than brown rice.
In practice, reaching a desirable intake of thiamin comes
usually from eating a wide variety of foods. Nineteenth-century
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Nutrition: Beriberi, A Deficiency Related to Grains
Table 1 Thiamin contributed by the different parts of a sample of dehusked rice grains
Dissected parts of rice grain Proportion by weight (%)
Pericarp testa aleurone layers 6 3.1
Germ area
Embryo 1 5.9
Scutellum 1 18.9
Endosperm 92
Total 100
Reproduced with permission from Carpenter KJ (2000) Beriberi, White Rice and Vitamin B. Berkeley, CA: University of California Press.
Japanese believed that consumptions of most foods other than
rice during the summer months caused heat. Hence, food
consumption was almost entirely restricted to rice conse-
quently leading to an increase in beriberi during these months.
Is there an Antithiamin Problem?
There are many references in the literature to at least a suspi-
cion that certain foods and beverages may be responsible for
beriberi appearing in people whose intake of thiamin would
otherwise be adequate.
Thiaminases
Many species of fish contain enzymes in their viscera that split
thiamin molecules at the junction between its two ring struc-
tures. This was discovered when foxes, being reared for their fur
and fed on a mix containing a large proportion of whole raw
fish, developed a form of paralysis that responded to injections
with thiamin. A similar condition was seen later in cats that
had been fed on a canned food containing a large proportion
of whole fish. It was believed that the thiamin in the mix had
been largely destroyed after the mix had been prepared and was
waiting to be autoclaved.
These experiences led to investigations as to whether
humans could be similarly at risk, but it appears not. The
enzymes are not present in fish muscles (i.e., fillets), and
even where small fish are eaten whole, they are not ground
up with other items of diet before being cooked. Lastly, it was
found in animal studies that a subsequent meal with different
constituents was not affected by thiaminases being present in
an earlier meal.
Another source of thiaminases was found to be bracken,
and their presence explained the condition known as stag-
gers that occurs in horses that have been feeding on bracken.
Cooked bracken, in which the thiaminase was inactivated,
proved harmless to horses. The only known case of thiaminase
poisoning in humans occurred in a group exploring the inte-
rior of Australia in the 1860s. Running out of provisions on
their return journey, they lived on the sporocarps in the fronds
of a particular fern that is now known to contain a high level of
a particularly heat-resistant thiaminase. All four men devel-
oped leg weakness and lassitude; three died, and the survivor
remained lame even after his safe return.
81
Thiamin
Concentration in fraction (mg per 100 g) Contribution to 100 g grain (mg)
0.186

0:059
0:248
0:189
0.05 0.046
0.480
Heat-Stable Antithiamins?
It has been found that when the thiamin in a food comes into
contact with polyphenols such as caffeic acid, it no longer gives
a fluorescent product in the usual thiochrome procedure for
the estimation of thiamin. This led some workers to suppose
that drinking large quantities of tea or coffee, or chewing betel
nut all sources of polyphenols might induce a condition of
thiamin deficiency. However, biological assays have indicated
that the vitamin is still fully available.
When thiamin is incubated with garlic extracts, it undergoes
a reaction with the allicin present in which the thiazole ring
opens, and the sulfur atom in the ring links to the alkyl sulfide
to form a disulfide compound. This is not measured in the
thiochrome reaction, but in the body it is reduced to re-form
the active vitamin. In fact, compounds of this type can be
absorbed more efficiently by alcohol-damaged intestinal
walls than ordinary thiamin. Thiamin tetra-hydrofurfuryl
disulfide in particular is approved for this purpose in some
countries.
Although there is no confirmed evidence of naturally occur-
ring heat-stable compounds that inactivate thiamin, chemists
have synthesized such materials. One, named oxythiamin,
has the amino group attached to the pyrimidine group in
thiamin replaced by a hydroxy group. Giving it to animals
results in the more rapid production of some of the signs of
thiamin deficiency, although it differs from thiamin in being
unable to pass the bloodbrain barrier.
Acute Deficiency in the West
With the discovery that autoclaving yeast powder would
destroy the thiamin, whereas the other B-vitamins were
retained, it was possible to place volunteers on an artificial
diet essentially free of thiamin. To the surprise of investigators,
some subjects had lost appetite within 2 weeks and became
nauseated and dizzy, with other mental symptoms, by 6 weeks,
but with no sign of peripheral nerve damage or cardiac abnor-
mality, which are characteristic of classic beriberi.
Trials using pigeons and rats with very deficient diets also
produced appetite loss and death before any sign of leg weak-
ness had developed. It appeared that in both humans and
animals, acute deficiency of thiamin resulted in damage to
the central nervous system. With slightly higher intakes, the
82 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Nutrition: Beriberi, A Deficiency Related to Grains
CNS had priority, whereas peripheral nerves gradually
degenerated.
Deficiencies in Total Parenteral Nutrition
There are many reports of people recovering from surgery of
the gastrointestinal tract who have developed acute thiamin
deficiency. They had been fed intravenously with a solution
providing energy, amino acids, and minerals, but no vitamins.
This is adequate for a short period, but thiamin is the first
vitamin to become depleted.
In a number of cases where this type of parenteral feeding
has continued for some weeks, a condition called Wernickes
encephalopathy has developed. Patients are confused and
have characteristic involuntary eye movements. Where patients
have died, autopsies have shown brain lesions analogous to
those found in acutely deficient animals. The same outcome
has been seen in subjects voluntarily fasting for long periods or
being unable to take food because of persistent vomiting in
pregnancy.
Alcoholism
One material that can be responsible for the production of
thiamin deficiency is ethanol (i.e., alcohol in everyday
speech). In developed countries where nearly everyone can
afford a well-balanced diet, most of those diagnosed as being
thiamin-deficient are alcoholics. The continued ingestion of
high levels of alcoholic beverages has many undesirable effects.
In the present context, two are relevant. First, the alcoholic
typically no longer bothers to eat a normal range of foods,
partly because the beverages provide a large portion of their
calorie needs and partly from nothing but their next drink
being of immediate interest. Second, the high level of alcohol
ingestion damages the intestinal wall so that thiamin is
absorbed less efficiently, and the requirement for the vitamin
increases.
Unfortunately, a small proportion of such victims develop
Wernickes encephalopathy, which may lead in turn to
Korsakoffs psychosis. Such people, sometimes referred to as
suffering from the WernickeKorsakoff syndrome, are at pre-
sent incurable and have to be maintained in a mental hospital
for the rest of their life. The cost of this to the state is such that
some specialists have argued that it would actually be cheaper
to have all beer and wine fortified with thiamin as a preventive.
There may be a genetic factor making some Western people
susceptible to the cerebral form of beriberi and the Wernick-
eKorsakoff syndrome, since it was seen even in Western pris-
oners of the Japanese in the Second World War who had white
rice but no alcohol, but apparently has not been seen in Asian
subjects.
Exercises for Revision
In your community, what dietary habits are most likely to
result in insufficient thiamin consumption?
What are the mandatory (and common voluntary) enrich-
ments of cereal grain products in your country? Do you
think the regulations make sense in relation to current
typical dietary patterns?
Exercises for Readers to Explore the Topic Further
What methods are used to measure thiamin content? How
stable is thiamin after typical food processing operations?
Are there any potential genetic solutions to low thiamin
content in milled rice?
See also: Food Grains and the Consumer: Consumer Trends in
Grain Consumption; Cultural Differences in Processing and
Consumption; Grain Composition and Analysis: The Composition
of Food Grains and Grain-Based Products; Non-wheat Foods: Rice:
Chinese Food Uses; The Basics: Grain and Plant Morphology of
Cereals and how characters can be used to identify varieties; Grain:
Morphology of Internal Structure.
Further Reading
Carpenter KJ (2000) Beriberi, White Rice and Vitamin B. Berkeley, CA: University of
California Press.
Cook CC, Hallwood PM, and Thomson AD (1998) B vitamin deficiency and
neuropsychiatric syndromes in alcohol misuse. Alcohol and Alcoholism
33: 317336.
Eijkman C (1929) Antineuritic vitamin and beriberi. In: Nobel Lectures: Physiology or
Medicine, pp. 19221941. Amsterdam: Elsevier.
Hinton JJC (1948) The distribution of vitamin B1 in the rice grain. British Journal of
Nutrition 2: 237241.
Jansen BCP and Donath WF (1926) On the isolation of the anti-beriberi vitamin.
Koninklijke Akademie von Wetenschappen, Amsterdam, Proceedings
29: 13901400.
Sauberlich HE, Herman YF, Stevens CO, and Herman RH (1979) The thiamin
requirement of the adult human. American Journal of Clinical Nutrition
32: 22372248.
Shimazono N and Katsura E (1965) Review of Japanese Literature on Beriberi and
Thiamine. Kyoto: Vitamin B Research Committee of Japan.
Takaki K (1906) The preservation of health amongst the personnel of the Japanese army
and navy. Lancet i: 13691374.
Truswell AS and Apeagyei F (1982) Alcohol and cerebral thiamin deficiency.
In: Jelliffe EF and Jelliffe DB (eds.) Adverse Effects of Foods, pp. 253258.
New York: Plenum Press.
Williams RR (1961) Toward the Conquest of Beriberi. Cambridge, MA: Harvard
University Press.
Williams RR and Cline JK (1936) Synthesis of vitamin B1. Journal of the American
Chemical Society 58: 15041505.
Wuest HM (1962) The history of thiamine. Annals of the New York Academy of Sciences
98: 385400.
 Celiac Disease
P Koehler, H Wieser, and KA Scherf, Deutsche Forschungsanstalt fur Lebensmittelchemie, Leibniz Institut, Freising, Germany
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Celiac disease (CD) is an inflammatory disease of the upper
small intestine triggered by dietary gluten from wheat, rye,
barley, and oats and has a prevalence of 1:300 to 1:100
individuals.
CD patients present with intra- and/or extraintestinal symp-
toms with villous atrophy as the classical clinical feature.
Gluten proteins are resistant to gastrointestinal digestion
and activate the adaptive and innate immune responses in
the lamina propria.
Other gluten-dependent diseases are dermatitis herpetifor-
mis, gluten ataxia, wheat-dependent exercise-induced
anaphylaxis, and non-celiac gluten sensitivity.
CD diagnosis is based on symptoms, serology, small intes-
tinal histology, and response to a gluten-free diet, which is
the current essential treatment.
Testing for CD toxicity may be done by in vivo challenge of
CD patients, by in vitro tests with tissues and cells of CD
patients, or with animal models.
Learning Objective
Attain understanding of symptoms, epidemiology, genetics,
pathogenesis, diagnosis, and therapy of celiac disease and
of its precipitating factor, gluten.
Introduction
Celiac disease (CD) is a permanent intestinal intolerance to
dietary wheat, rye, barley, and oat proteins (gluten) that pro-
duce mucosal lesions and nutrient malabsorption in geneti-
cally susceptible individuals. The current essential therapy is a
strict lifelong withdrawal of gluten from the diet. The preva-
lence of CD was underestimated for a long time; with the
development of sensitive serological tests, more and more
evidence is emerging that CD is one of the most frequent
food intolerances in many parts of the world. An overview of
extensive research on CD is provided.
Definition
CD, also known as celiac sprue or gluten-sensitive enteropathy,
may be defined as an inflammatory disease of the upper small
intestine (duodenum and jejunum) in genetically susceptible
individuals. CD is a permanent food intolerance induced by
ingestion of storage proteins (gluten) from certain cereals
(wheat, rye, barley, and oats) that cause damage to the enter-
ocytes and, as a consequence, malabsorption of important
nutrients. CD patients develop a flat jejunal mucosa with the
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00083-8
absence of normal villi, a cellular infiltrate of the lamina
propria, and an increase of the number of intraepithelial lym-
phocytes (IELs). The mucosa improves morphologically on
treatment with a gluten-free diet (GFD) and relapses when
gluten is reintroduced. The clinical features of CD are highly
variable and can be divided into intestinal symptoms (e.g.,
diarrhea, steatorrhea, and vomiting) and extraintestinal com-
plaints (e.g., osteoporosis, anemia, and constant fatigue). Ther-
apeutically, a lifelong GFD is necessary.
Historical Review
CD was already noted by the Roman physician Aretaeus of
Cappadocia as early as the second century AD, who wrote
that if diarrhea does not proceed from a slight cause of only
one or two days duration and if, in addition, the general system
be debilitated by atrophy of the body, the celiac sprue of
chronic nature is formed. However, it was not until 1888
that the English physician Samuel Gee described the classical
account of CD features. He recognized that if the patient can
be cured at all, it must be by means of the diet. Thereupon, the
dietary treatment of CD was continued more or less success-
fully. For example, all sources of carbohydrates such as bread,
cereals, and potatoes were excluded or a banana diet was
recommended. Such extreme dietary treatments were then
used for many years.
In 1950, Willem K. Dicke, a Dutch pediatrician, observed a
decline in CD in the Netherlands during the grain-deprived
years of the Second World War. A clear association between CD
and the ingestion of wheat and, later on, of rye and barley was
established. The fractionation of wheat flour and testing led to
the conclusion that gluten was toxic, whereas starch and the
water-soluble albumins were not. Since that time, a GFD has
been the conventional treatment of CD, which excludes prod-
ucts containing wheat, rye, and barley, only excepting pure
starches produced from these cereals. The celiac toxicity of
oats has been judged controversially. With the introduction
of peroral biopsy of the intestinal mucosa, the abnormalities
of the small intestinal mucosa tissue of CD patients were
demonstrated, and among others, this diagnostic procedure is
the criterion standard of diagnosis to this day.
Symptoms and Pathology
A range of symptoms may be associated with CD, and these
can be divided into intestinal disorders and extraintestinal
features caused by malabsorption of essential nutrients such
as deficiencies of vitamins and minerals. In infants, classical
symptoms appear after weaning and introduction of cereals
into the diet: diarrhea, abdominal distension, failure to thrive,
vomiting, muscle wasting, and apathy. Older children tend to
83
84 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease
have more varied symptoms. Besides diarrhea or constipation,
features such as anemia, loss of appetite, and short stature may
be predominant. Diarrhea is the main presenting feature of
adults; in some patients, anemia, osteoporosis, abdominal
pain, loss of weight, and weakness may be found. A minor
number of patients present psychological or psychiatric symp-
toms such as depression, epilepsy, autism, or schizophrenia.
Frequently, the disease may be clinically silent or masked by
associated diseases like type 1 diabetes, although the patients
exhibit the full mucosal lesion. They are discovered only when
intestinal studies are undertaken. Around 25% of CD patients
develop a resistance to a strict GFD. This condition, called
refractory celiac disease (RCD), is characterized by a constant
inflammatory process in the intestinal mucosa accompanied
by persistent diarrhea, abdominal pain, and weight loss. RCD
carries a significant burden of morbidity and mortality, mainly
from malnutrition and lymphoma.
The small intestinal lesion of CD has a highly characteristic
morphology, but it may vary from patient to patient depending
on the severity and extent of disease. Patients show a wide
variation of mucosa appearance from a complete flat to a
convoluted pattern. The parameters usually assessed for the
measurement of intestinal mucosa are villous height, villous
width, villous height to crypt depth ratio, mucosal thickness,
epithelial cell height, crypt mitotic activity, and number of
IELs. Electron microscopy demonstrates fewer microvillous
intramembrane particles and abnormal tight junctions (TJs)
between epithelial cells. In untreated CD, serological anti-
bodies to gliadin, deamidated gliadin peptides, reticulin, endo-
mysium, and tissue transglutaminase (TG2) are present.
Epidemiology and Genetics
CD is prevalent mostly in Europe and those countries to which
Europeans have emigrated including North and South America
and Australia. However, CD is also known to occur in many
other parts of the world. In the past, CD was considered a
comparatively uncommon disorder with prevalence rates of 1
in 2000 to 1 in 1000. Several recent studies using serological
screening followed by small intestinal biopsy have shown a
much higher prevalence, and it is now estimated that CD may
affect as many as 1 in 300 to 1 in 100 individuals. The iceberg is
a common model to describe the epidemiology of CD. The tip
of the iceberg is formed by patients who have been diagnosed
by conducting a biopsy demonstrating a flat mucosa. Below
the waterline, there is a big group of undiagnosed patients with
a flat mucosa, but with no or weak symptoms (undiagnosed
silent CD). At the bottom of the iceberg, there is a small section
of patients with a normal mucosa but with the genetic predis-
position and increased levels of celiac-specific antibodies.
These subjects may develop clinically overt CD later in life
(undiagnosed potential CD).
The occurrence of CD among first-degree relatives of CD
patients is fairly strong with a prevalence of 1015%. The
concordance between dizygotic twins is reported to be in a
range of 1120% and that between monozygotic twins
7580%. CD can occur at any age, but in adults, the peak
incidence is in the fifth decade. Females are more commonly
affected than males with a suggested ratio of around 2:1.
CD is associated with histocompatibility complex class II
alleles HLA-DQ2 and HLA-DQ8. Almost all CD patients
(
97%) possess one of these HLA genes. The large majority
of patients (
9095%) are DQ2-positive, and the remainder
are DQ8-positive. Two isoforms DQ2.5 (predominant) and
DQ2.2 (rare) were identified among DQ2 alleles. HLA-DQ
genes encode protein heterodimers with a- and b-chains,
both responsible for the CD-specific binding of gluten peptides
by intestinal antigen-presenting cells. The HLA-DQ2.5 geno-
type is associated with a very high risk for developing CD,
followed by DQ8 (high) and DQ2.2 (low). However, about
30% of the healthy population carry DQ2 and will never
develop CD. Thus, HLA-DQ genes alone do not explain the
development of CD, and additional factors such as non-HLA
genes and environmental factors need to be involved in disease
development. Further, CD-specific genome research, mainly
focused on non-HLA genes, indicated little consensus, and
the contribution of all non-HLA genes, known up to now,
has been estimated to account for less than 10%.
Different environmental factors have been considered to be
important for the development of CD. For example, infections
by a variety of pathogens, an imbalanced intestinal microbiota,
the amount and timing of gluten exposure, and breastfeeding
during first gluten intake have been associated with developing
CD in infants.
Pathogenesis
The development of CD is induced by the oral intake of gluten
present in wheat, rye, barley, and oat products. A daily intake
of more than 20 mg of gluten is estimated to be not tolerable
for CD patients. This approximately correspond to one-
hundredth of a slice of bread. In contrast to most other food
proteins, gluten proteins are highly resistant to complete pro-
teolytic digestion within the gastrointestinal tract due to their
high proline content. Thus, relatively large protein fragments
(peptides) accumulate in the small intestine and reach the
intestinal brush border (step 1 in Figure 1). However, the
poor digestibility of gluten proteins is not sufficient to cause
CD, and there is no known difference between healthy indi-
viduals and CD patients in their (in)ability to digest these
proteins. Changes in the mucosal permeability for gluten pep-
tides are supposed to be the first pathogenic event in the
development of CD. Gluten peptides are bound to the intesti-
nal mucosa before passing through the enterocyte layer either
transcellularly or paracellularly (Figure 1, step 2). The most
important structures of the paracellular route are the epithelial
pore-forming TJs. They are composed of a number of proteins
(e.g., zonulins, claudins, and occludin) that regulate paracel-
lular permeability. Zonulin 1, known to enhance permeability
and to be upregulated during the acute phase of CD, has been
proposed to be one of the initial factors of the pathogenesis of
CD. When crossing the epithelium via the transcellular path-
way, gluten peptides are differently degraded in the intracellu-
lar compartments of the enterocytes of CD patients. The
transcellular permeability is much higher for intact CD-active
peptides compared with nonactive peptides.
After passing through the epithelium, CD-active peptides
arrive at the lamina propria and induce two different immune
responses: the adaptive and the innate responses. Within the
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease 85

I
1

E E 2 E E
II

IFN
5 TNF
T
MMP A B
III P T IgA
C B P
T IgG
4 B 6
T
B
Th 1
Th 2

gluten peptide deamidated gluten peptide TG2 TG2/gluten peptide conjugate

TG2/deamidated gluten peptide conjugate
Figure 1 Schematic representation of the adaptive immune response in the intestinal lymphatic tissue and destruction of enterocytes in the
pathogenesis of celiac disease. I, intestinal lumen; II, epithelium; III, lamina propria; APC, antigen-presenting cell; B, B cell; E, enterocyte; IFN, interferon-
g; Ig, immunoglobulin; MMP, matrix metalloproteinase; P, plasma cell; T, CD4 T cell; TG2, tissue transglutaminase; Th, T helper; TNF, tumor
necrosis factor. For steps 16, see text. Reproduced with permission from Wieser H, Koehler P, and Konitzer K (2014) Celiac Disease and Gluten:
Multidisciplinary Challenges and Opportunities, 1st edn. London, Waltham, San Diego: Academic Press.

adaptive mechanism, gluten peptides are specifically modified
by TG2 (Figure 1, step 3) either by partial deamidation of
glutamine residues leading to negatively charged glutamic acid
residues or by cross-linking to TG2 by isopeptide bonds between
glutamine residues of the peptides and lysine residues of TG2.
Particularly, deamidated peptides are selectively bound to DQ2/
DQ8 heterodimers on the surface of antigen-presenting cells
(dendritic cells, macrophages, and B cells) and then presented
to the T cell receptor of gluten-sensitive CD4 T cells. The stim-
ulation of T cells leads to the secretion of proinflammatory T
helper (Th-1) cytokines (Figure 1, step 5), in particular
interferon-g, tumor necrosis factor, matrix metalloproteinase,
and nitric oxide, which lead to premature senescence of small
intestinal enterocytes, shedding of these cells, and, in turn, the
observed small intestinal enteropathy. Simultaneously, T cells
provide help for the production of IgA and IgG antibodies to
gluten proteins and autoantibodies to TG2 (Figure 1, step 6).
In contrast, the rapid innate immune system is characterized
by the massive increase in IELs, one of the hallmarks of CD, which
can be observed very early before the onset of villous atrophy. Two
subsets of IELs bearing the abT cell receptor or the gdT cell
receptor are linked to CD innate immunity. Interleukin-15
(IL-15) has been considered a central player in this part of the
gluten-induced immune response. IL-15 is produced by epithelial
and lamina propria cells in active CD, but not by T and B cells
involved in the adaptive immune response. Recently, it has been
suggested that amylase-trypsin inhibitors (ATIs) present in cereal
flours are coactivators of the innate immune system. How the
cross talk between innate and adaptive immunities is orchestrated
is an outstanding question that will need to be addressed.
Associated Diseases
Dermatitis herpetiformis, a skin disorder, is associated with
CD, because some degree of gluten-sensitive enteropathy is
common to both conditions. It appears as patches of itchy
red papules and blisters on the extensor surface and pressure
areas; healing up results in pigmentation. Histologically, there
is infiltration of the dermal papillae with inflammatory cells.
The dermal histology is similar to that of the small intestine in
CD, but the abnormalities tend to be milder and patchier. In
many cases, a GFD results in a significant improvement.
Gluten ataxia is one of the neurological manifestations
attributed to gluten intake. All these patients present gait
ataxia, and the majority have limb ataxia and evidence of
cerebellar atrophy. The current diagnostic recommendation
is a serological screening for antigliadin, anti-TG2, and anti-
TG6 antibodies. Patients positive for these antibodies should
follow a GFD.
Wheat-dependent exercise-induced anaphylaxis (WDEIA) is
a severe IgE-mediated allergic reaction provoked by the com-
bination of wheat ingestion with intense physical exercise over
the next few hours. Aspirin and ethanol, together with wheat-
containing food, are further cofactors for WDEIA. Peptides
derived from o5-gliadins and high-molecular-weight (HMW)-
glutenin subunits have been identified to be critical for IgE
binding.
Non-celiac gluten sensitivity (NCGS) is a gluten-mediated
disorder characterized by gastrointestinal complaints including
bloating, diarrhea, weight loss, and abdominal pain. The small
intestine of patients is usually normal, and serum antibodies
typical for CD are absent. ATI or fermentable oligo-, di-, and
monosaccharides and polyols may be implicated in NCGS, but
to date, there is too little knowledge on responsible triggers of
this currently underrecognized and undertreated disorder.
Recent screening studies have shown an increased preva-
lence of CD in autoimmune disorders such as type 1 diabetes,
thyroid disease, primary biliary cirrhosis, and Sjogrens syn-
drome. Moreover, IgA deficiency, chronic fibrosin alveolitis,
other interstitial lung diseases, and concomitant distal ulcera-
tive colitis have been reported in association with CD. In other
86 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease
instances, there are unexpected associations such as epilepsy
and various undefined neurological disorders. A small number
of patients with CD have exhibited symptoms of various psy-
chiatric disorders, such as schizophrenia. A lymphoma of the
small intestine is the classical malignancy associated with CD.
The prevalence of carcinoma of the small intestine, esophagus,
or pharynx, crypt hyperplasia, villous atrophy, and mucosal
hypertrophy also increases with CD. It has been reported that a
strict GFD helps safeguard against these malignancies.
Diagnosis
The diagnosis of CD is complex and tends to require a high
level of clinical suspicion. The diagnostic approaches have
changed during the last decades, and recommendations for
the procedures are still different. In summary, the diagnostic
scheme may consist of the following points: (1) clinical history
and symptomatology, (2) serology, (3) small intestinal
histology, (4) response to a GFD, and (5) HLA status. Classical
intestinal symptoms should prompt the primary doctor to
initiate a serological test. Extraintestinal symptoms are more
difficult to interpret, and medical specialists (e.g., dermatolo-
gists, dentists, and neurologists) should be aware of possible
CD. For initial testing, a quantitative test detecting IgA class
antiendomysium, anti-TG2, and antideamidated gliadin pep-
tide antibodies from a blood sample is recommended. To
achieve the clinical demand for both maximal sensitivity and
specificity, the combination of different tests is recommended.
In case of IgA deficiency, IgG class antibodies have to be used.
The histological judgment of small intestinal mucosa is
commonly regarded as the gold standard for the reliable diag-
nosis of CD. Biopsy specimens of active CD demonstrate a flat
mucosa with the absence of intestinal villi, a cellular infiltrate
of the lamina propria, and an increase in the number of IELs.
In the ESPGHAN guidelines revised in 2012, the omission of
the small intestinal biopsy is only tolerated in children and
adolescents with distinct signs and symptoms suggestive for
CD and very high anti-TG2 antibody titers with levels exceed-
ing ten times the upper limit of normal.
Genetic tests based on HLA-DQ2 and DQ8 alleles cannot
be used to confirm CD, because they are common in the
general population (around 30%). In the case of doubtful
diagnostic results, however, HLA-DQ genotyping can be used
to rule out the existence of CD, because of its high negative
predictive value. If a patient is negative for HLA-DQ2/8, CD
can be excluded lifelong and no further investigations and
follow-up are needed. The final confirmation of CD is estab-
lished by a clinical, serological, and histological response to a
GFD. When keeping a strict GFD, patients show a rapid
improvement of symptoms within weeks. Histological regen-
eration is slower and complete recovery can take months or
years.
Therapy
After the diagnosis of CD has been established, permanent
withdrawal of gluten from the diet is the current essential
treatment. The GFD involves avoiding products containing
wheat, rye, and barley. Starches of these cereals are permitted,
if the gluten level is lower than 20 mg kg1 (Draft Revised
Codex Standard 2008). The toxicity of oats has been discussed
controversially, but recent studies have provided strong evi-
dence that only a very small subgroup of CD patients cannot
tolerate oats. The daily intake of gluten should not exceed
20 mg. Since small amounts of gluten are hidden in many
foods, dietary counseling is absolutely necessary. In most
countries, celiac societies are an invaluable help in providing
dietary guidelines and food lists. The prognosis for patients
who are correctly treated is excellent, whereas failure to imple-
ment a strict diet may result in continuing symptoms and in
the two major complications: osteoporosis and malignancy.
CD patients may consume gluten-free foods from two differ-
ent categories. Firstly, they are allowed to eat a wide range of
common products such as meat, fish, milk, fruits, and vegeta-
bles. In the case of composite foods, however, it is difficult to
recognize whether they are gluten-free or not. Labeling of gluten
as an allergen is mandatory in the framework of CODEX Ali-
mentarius and in the European Union. Secondly, patients con-
sume dietetic food that is gluten-free according to the CODEX
Alimentarius standard. The current standard and legislation in
the United States, Canada, and the European Union set a gluten
level of 20 mg kg1 food as the threshold for a gluten-free claim.
In some countries, foods containing up to 100 mg gluten/kg,
such as wheat starch, can be labeled as low in gluten. For gluten
analysis, extraction with 60% ethanol and an immunochemical
method for quantitative determination (enzyme-linked immu-
nosorbent assay (ELISA) with monoclonal or polyclonal anti-
bodies) are recommended. Dietetic gluten-free foods are mostly
substitutes of products usually containing wheat, rye, and bar-
ley, such as bread, other baked products, pasta, and beer. Com-
mon raw materials used for these products are flours from corn,
sorghum, rice, buckwheat, quinoa, teff, and chestnut, and com-
mon thickening agents are locust bean gum, guaran gum, and
hydroxypropyl methyl cellulose.
Owing to the enormous increase of knowledge about the
pathomechanism of CD, a number of novel strategies for the
prevention and treatment of CD have been developed. Usually,
CD-toxic peptides derived from digested gluten survive the
normal digestive process. Supplementation with additional
peptidases may degrade CD-toxic epitopes to avert an immune
response. Peptidases from bacteria, fungi, and germinating
cereals have been proposed for oral therapy. Some of these
approaches are already in phase 1 and 2 clinical trials. More-
over, inhibitors of zonulin, a key molecule for intestinal
permeability; inhibitors of TG2, the autoimmunogen of CD;
inhibition of gluten peptide presentation by HLA-DQ2
antagonists; and modulation or inhibition of proinflammatory
cytokines have been proposed as alternative therapies. How-
ever, the risks, benefits, and costs of alternatives have to be
carefully weighed, and the conditions and indications under
which such alternative therapies might be warranted have to be
accurately defined.
Toxicity Testing
Numerous in vivo and in vitro methods have been used to
identify CD toxicity and immunogenicity of different cereal
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease
and noncereal materials, proteins, and peptides. The tests can
be differentiated into (1) in vivo challenge of patients, (2)
in vitro tests with tissues and cells of patients, and (3) animal
models. Most investigators would agree that in vivo testing is
the gold standard for assessing CD toxicity of proteins or
peptides. Early workers established toxicity in a series of feed-
ing tests based on the production of symptoms such as steat-
orrhea or on tests such as malabsorption of fat or xylose.
However, an important impediment was that CD patients
differed widely in their sensitivity to gluten and, consequently,
the optimal amount of gluten equivalent used to challenge
patients and the duration of challenge were uncertain. In any
case, 10100 g of gluten equivalent was necessary for the test-
ing of each patient, and such large amounts were the most
crucial limiting factor for feeding tests of purified proteins or
peptides. Direct instillation into the small intestine followed
by biopsy allows the reduction of required amounts to 1 g
equivalent of gluten. Biopsies are taken from patients in remis-
sion at the beginning and after several hours of testing.
Changes in enterocyte height, ratio of villous height to crypt
depth, and the number of IELs are considered to be reliable
parameters for toxicity assessment.
Because in vivo tests require large quantities of substances
and only a limited number of test patients are available, a series
of in vitro tests has been developed. The organ culture of
human small intestine, which requires only milligram equiva-
lents of gluten, provides the most reliable in vitro approach.
The intestinal tissue from patients with active CD is removed as
part of the diagnostic procedure and incubated in a culture
medium. The tissue shows improvement of enzyme activity
and morphology in the medium alone, but not in the presence
of CD-toxic substances. Apart from human material, tissue
cultures with fetal rat and chicken intestine have been used. T
cell lines and clones raised against gluten are useful for per-
forming a large number of experiments and readily demon-
strate the T cell response to stimuli. Gluten-sensitive T cells can
be isolated from intestinal biopsies cultured and stimulated
in vitro or can be found in the blood of treated CD patients after
oral gluten challenge. A frequently used test is the T cell prolif-
eration assay performed by incubation of intestinal CD-
sensitive T cells, the putative antigen ( 100200 mg ml1)
treated with TG2, antigen-presenting cells (e.g., B cells), and
tritiated thymidine. The proliferation of T cells determined by
scintillation measurements is a parameter for immunogenic
effects. Additionally, the production of IFN-g or interleukins
can be measured. T cell tests have been used widely to compare
the level of immunogenic effects. Generally, it should be men-
tioned that T cells do not necessarily reflect the small intestine
in vivo, because they lack any connection to other cell types
present in the intestine. In addition, T cells are frequently
different in their reaction to antigens; application of several
simultaneous T cell lines from different patients is therefore
needed. Moreover, immunogenicity does not always corre-
spond to toxicity demonstrated by in vivo or organ culture
tests. In any case, in vivo testing ultimately will be necessary
to evaluate conclusions on in vitro testing. Among animals,
there are three models that spontaneously generate gluten-
dependent diarrhea: the Irish setter dog, the rhesus macaque,
and the horse. However, the animal model that reproduces all
important aspects of CD still awaits development.
87
Toxicity of Cereals
The relationship between CD and the ingestion of wheat flour
was established in 1950. Soon after, a series of investigations
led to the conclusion that rye and barley were also harmful,
whereas corn, rice, buckwheat, and potatoes were not. There
has been disagreement about the toxicity of oats; the reason for
that could have been that commercial oat flour is frequently
contaminated with small amounts of wheat, rye, or barley.
Most clinical studies indicate that pure oats do not activate
CD and it is proposed that only a very small subgroup of CD
patients is sensitive to oats. Accordingly, CD is closely related
to the taxonomy of cereals: only species found in the tribe
Triticeae within the grass family Poaceae are likely to trigger
CD. All wheat species besides bread wheat such as durum
wheat, spelt, emmer, einkorn, and kamut have been suggested
to be toxic for CD patients due to homology of their storage
proteins to wheat.
Toxicity of Cereal Proteins
Immediately after wheat was established as the CD-activating
cereal, fractionation of the flour and in vivo testing led to the
conclusion that gluten is toxic, whereas starch and albumins
are not. Gluten is the rubbery protein mass that remains when
wheat dough is washed with water to remove starch granules
and other soluble constituents. Wheat gluten proteins can be
divided according to their solubility in aqueous alcohols into
the soluble gliadins and the insoluble glutenins. Toxicity tests
indicated that the gliadin fraction was the most toxic factor,
whereas the effect of the glutenin fraction was described either
nontoxic, weakly toxic, or as toxic as gliadins, but on very
inadequate evidence. More recent studies on pure glutenin
subunits indicated that the glutenin fraction is also harmful
for CD patients.
Flour protein fractions of rye and barley have not been
tested for celiac toxicity until now, but equivalent to the gliadin
fraction of wheat, the corresponding prolamin fractions of rye
(secalins) and barley (hordeins) are associated with CD.
Although the prolamin fractions of cereals are crude mixtures
of different proteins, their amino acid compositions (Table 1)
show a close relationship to both taxonomy and CD toxicity.
Toxic gliadin, secalin, and hordein fractions are characterized
by the highest contents of glutamine (3537 mol%) and pro-
line (1723 mol%). Both amino acids have been considered to
be important for CD toxicity. The prolamins of rice, millet, and
corn are lower in glutamine and proline, but rich in alanine
(914 mol%) and leucine (1219 mol%). Oats are in an inter-
mediate position, the glutamine content being similar to the
Triticeae prolamins and the values of proline and leucine in
agreement with those of other species.
Among the prolamin fractions, only wheat prolamin (glia-
din) has been investigated in detail. The digestion with pepsin
and trypsin alone or followed by pancreatin resulted in the
retention of toxicity, and a peptide mixture with a molecular
mass less than 2000 was still toxic. Also, the breakdown of
the disulfide bonds by oxidation or heating during the
baking process did not abolish toxicity. In consequence, the
88 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease

Table 1 Partial amino acid composition (mol%) of prolamins of different cereals

Amino acid Wheat (gliadin) Rye (secalin) Barley (hordein) Oats (avenin) Rice (oryzin) Millet (kafirin) Maize (zein)

Glx 37.1 35.4 35.3 34.1 19.6 21.8 19.4

Pro 16.6 18.4 23.0 10.2 5.1 7.8 10.2
Leu 6.9 5.8 6.1 10.6 11.8 13.4 18.5
Ala 2.8 3.0 2.3 5.5 9.1 13.5 13.6
Met 1.1 1.0 0.9 2.1 0.5 1.7 1.1
Lys 0.8 1.0 0.5 1.0 0.5 0.0 0.0
Trp 0.4 0.4 0.6 0.3 0.5 1.5 0.0

Table 2 Amino acid sequences of peptides tested for CD toxicity by instillation and organ culture tests

Amino acid sequence Source Position Toxicitya

VPVPQLQPQNPSQQQPQEQVPL Gliadin a324 (OC)

VQQQQFPGQQQPFPQQPYPQPQPFPSQQPY Gliadin a2555 (OC)
VRVPVPQLQPQNPSQQQPQEQVPLVQQQQF A-Gliadin a130 (OC)
LGQQQPFPPQQPYPQPQPFPSQQPY A-Gliadin a3155 (OC)
LGQGSFRPSQQN Synthetic a206217 (IN)
LGQQQPFPPQQPYPQPQPF Synthetic a3149 (IN, OC)
LGQQQPFPPQQPY Synthetic a3143 (IN)
PQPQPFPSQQPY Synthetic a4455 (IN)
LQLQPFPQPQLPYPQPQLPY Synthetic a5675 (IN)
SQQPYLQLQPFPQPQLPYSQ Synthetic a5170 (OC)
a
IN, instillation; OC, organ culture.

three-dimensional structure of gliadin proteins is not important Toxicity of Peptides

for the toxic effect. The complete degradation into free amino
acids by acid hydrolysis, however, rendered it harmless, as did
extensive deamidation of glutamine side chains with a limited
cleavage of peptide chains. Subsequent in vivo and in vitro studies
demonstrated that all gliadin types produce toxic effects. In
contrast, the major protein types of the glutenin fraction, low-
molecular-weight (LMW)-glutenin subunits have not been
tested in vivo until now. However, in vivo studies with HMW
glutenin subunits confirmed their toxicity for CD patients.
Currently, approaches aimed at reducing or abolishing CD
toxicity of wheat are underway. Beside the strategy to reduce
the toxicity of wheat by conventional breeding approaches, the
recombinant production of completely safe gluten proteins is
an additional aim based on currently gained knowledge on the
elimination of the toxic fragments. The first approach is based
on the assumption that the dose of exposure to gluten-derived
CD-active epitopes will contribute to a general reduction of the
prevalence of CD and of symptom severity in the population.
Barley is a diploid cereal with pure inbred cultivars available
and is, therefore, attractive for use in genetic approaches to
detoxification. Barley lines lacking B-, C-, and D-hordeins gave
gluten concentrations of 45 mg kg1 after quantitation by
ELISA and liquid chromatography-mass spectrometry (LC-MS).
Test with peripheral blood mononuclear cells activated by
oral hordein challenge demonstrated that the immunogenicity
of these lines was reduced 20-fold compared with that of
wild-type barley. Therefore, the creation of hordein-free barley
seems to be a realistic goal. However, the quality of products
(e.g., beer) derived from hordein-free barley lines has still to be
demonstrated.
The discovery that gliadins are enzymatically degraded without
loss of toxicity and subsequent studies on gluten digestion and
immune response clearly revealed that gluten peptides are, in
fact, the precipitating factors of CD. Therefore, research on CD
toxicity has been focused on peptides either isolated from
protein digests or synthesized. In vivo instillation tests and
in vitro organ cultures enabled the study on peptides in milli-
gram and even microgram amounts. With the exception of
one peptide (a206217), toxic peptides are contained in the
N-terminal domain of a-gliadins characterized by sequences
rich in glutamine (Q) and proline (P) (Table 2). Since the end
of the 1990s, the search for CD-active peptides has shifted
toward epitopes that stimulate intestinal gluten-sensitive
T cell lines of CD patients, and peptides from gliadins, glute-
nins, secalins, and hordeins were investigated. A small selection
of the first generation (19992005) of immunogenic peptides
is presented in Table 3. Given the heterogeneity of wheat, rye,
and barley proteins, it is not surprising that a large number of T
cell epitopes exist. The major part of peptides is HLA-DQ2
restricted and is derived from the glutamine- and proline-rich
sequence domains. A number of studies have shown that
patients respond heterogeneously to different epitopes and
that there also are differences between children and adults.
The Food Allergy Research and Resource Program of the
Department of Food Science and Technology, University of
NebraskaLincoln, the United States, has compiled a database
of CD-active gluten peptides. It includes more than 1000
native or deamidated peptides described in more than 60
publications. These peptides have their origins in all types of
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease 89

Table 3 Amino acid sequences of selected gluten peptides would be helpful to establish a more precise epidemiological
stimulatory for intestinal T cells of CD patients picture of CD. Little is known on the complex interplay
between genetic and environmental factors leading to CD
Protein Amino acid sequencea
development, and it will be a major area of research to under-
a2 LQLQPFPQPQLPYPQPQLPYPQPQLPYPQPQPF stand their respective contributions and develop possible pre-
a20 PFRPQQPYPQPQPQ vention strategies. Further studies are required to allow a better
g5 FLQPQQPFPQQPQQPYPQQPQQPFPQ diagnostic differentiation between CD, NCGS, and irritable
g5 QFSQPQQQFPQPQ bowel syndrome, because the symptoms and nonspecific find-
LMW 156 QQQQPPFSQQQQSPFSQQQQ ings may be similar in patients. While the role of gluten pep-
LMW 17 QQPPFSQQQQQPLPQ tides in the adaptive immune response is well characterized,
HMW 2 GQQGYYPTSPQQS much less is known regarding the innate immune response.
HMW 2 QGYYPTSPQQSG
Elucidating the mechanism of the innate immune response
Sec a2 QPFPQPQQPFPQSQ
will contribute to a better understanding of the CD pathome-
Sec a9 PQQPFPQPQQPFPQ
Hor a2 QQFPQPQQPFPQQP chanism. Gluten-sensitive T cell assays are used to compare the
Hor a9 PQQPFPQPQQPFRQ level of immunogenic effects, but immunogenicity measured
Av a9A QYQPYPEQQEPFVQ by T cell tests does not necessarily correspond to toxicity
Av a9B QYQPYPEQQQPFVQ revealed by in vivo or organ culture tests. This is the reason
why it would be of interest to test immunogenic peptides by
a
Glutamine (Q) residues targeted by TG2 are in bold. in vivo assays. The GFD will remain the preferred therapy of CD
Source: Wieser H, Koehler P, and Konitzer K (2014) Celiac Disease and Gluten:
in the future. Regarding alternative therapies, CD patients have
Multidisciplinary Challenges and Opportunities, 1st edn. London, Waltham, San Diego:
expressed the wish to have access to a vaccine against CD or to
Academic Press, with permission.
a pill that would occasionally allow them to eat gluten-
containing food. Such alternative therapies are still in the
wheat gliadins and glutenins, rye secalins, barley hordeins, and
early stages of development, and they will have to undergo
oat avenins. Some authors have subdivided CD-active peptides
safety and toxicology testing before they can enter the market.
into toxic peptides generating an innate immune response
and immunogenic peptides generating an adaptive immune
response. The discrepancy between the numbers of immuno-
Exercises for Revision
genic peptides identified by T cell tests and of toxic peptides
identified by in vivo or organ culture tests is extraordinary: 1014
peptides have been classified as immunogenic, 10 peptides as
What is the primary environmental trigger for CD and
which cereals are harmful?
toxic, and 5 peptides as both toxic and immunogenic. The
question of whether immunogenicity corresponds to toxicity
What other gluten-related disorders are there?
in any case remains to be answered.
What are the steps in the diagnostic scheme to identify CD
and what is the essential treatment once the diagnosis has
In summary, a common feature among toxic/immunogenic
been established?
peptides is the presence of multiple proline and glutamine
residues, which gives rise to four unique structural and func-
What methods can be used to measure CD activity?
tional properties:
What foods are allowed to be consumed by CD patients?
What are the characteristic properties of CD-toxic/
1. These peptides are exceptionally resistant to peptidolysis immunogenic peptides?
by gastric, pancreatic, and intestinal digestive peptidases
because of their high proline content. As a result, a high
intestinal concentration of potentially immunogenic pep- Exercises for Readers to Explore the Topic Further
tides is maintained following a gluten-containing diet.
2. Selected glutamine residues in these gluten peptides are CD patients need to adhere to a strict GFD. Compare the
deamidated or transamidated by TG2 under physiological characteristics of wheat bread to those of gluten-free breads.
conditions, leading to enhanced immunogenicity. The increased knowledge about the pathomechanism of
3. The proline-rich gluten peptides naturally adopt a left- CD has led to the development of novel treatment strate-
handed polyproline II helical conformation, which is the gies, such as oral peptidase therapy. Look at the different
preferred conformation of all bound major histocompati- proposed treatment options and identify which steps in the
bility complex class II ligands. pathomechanism are targeted.
4. Large peptides that contain multiple HLA-DQ-binding epi- Take a look at the database of CD-active gluten peptides of
topes have a greater T cell stimulatory activity than small the Food Allergy Research and Resource Program of the
peptides containing only one epitope. Department of Food Science and Technology, University
of NebraskaLincoln, the United States (www.allergenon-
line.org/celiacbrowse.shtml), and find the CD-active pep-
tides derived from alpha-gliadins.
Future Prospects

A standardization of criteria for prevalence determination and See also: Appendix 1: Nutrient-Composition Tables for Grains and
a differentiation into cases of active, silent, and potential CD for Grain-Based Products; Appendix 2: Lists of Standardized Testing
90 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | Celiac Disease
Methods for the Analysis of Grain and Grain-Based Foods; Food
Grains: Intolerance, Allergy and Diseases: Cereal Allergens; The
Gluten-Free Diet; Grain Composition and Analysis: Standardized
Test Methods for Grains and Grain-Based Products; The Composition
of Food Grains and Grain-Based Products; Units of Grain Science and
Trade: Equivalence between the US, Chinese, and Metric Units;
Proteins: Protein Synthesis and Deposition; Proteomic Analysis
Relevant to Cereal Grains; The Protein Chemistry of Cereal Grains; The
Basics: Taxonomic Classification of Grain Species; The Grain Crops:
An Overview; ; The Cereal Grains: An Overview of the Family of
Cereal Grains Prominent in World Agriculture; Barley: An Overview of a
Versatile Cereal Grain with Many Food and Feed Uses; Oats: Overview;
Wheat: An Overview of the Grain That Provides Our Daily Bread;
Wheat Processing: Gluten and Modified Gluten; The Gluten Proteins
of the Wheat Grain in Relation to Flour Quality.
Further Reading
Abadie V, Sollid LM, Barreiro LB, and Jabri B (2011) Integration of genetic and
immunological insights into a model of celiac disease pathogenesis. Annual Review
of Immunology 29: 493525.
Arendt EK and Dal Bello F (2008) Gluten-Free Cereal Products and Beverages, 1st edn.
Burlington, San Diego, London: Academic Press.
Catassi C, Fabiani E, Iacono G, DAgate C, Francavilla R, Biagi F, et al. (2007) A
prospective, double-blind, placebo-controlled trial to establish a safe gluten
threshold for patients with celiac disease. American Journal of Clinical Nutrition
85: 160166.
Dicke WK, Weijers HA, and van de Kamer JH (1953) Coeliac disease. II. The presence in
wheat of a factor having a deleterious effect in cases of coeliac disease. Acta
Paediatrica 42: 3442.
Husby S, Koletzko S, Korponay-Szabo IR, Mearin ML, Phillips A, Shamir R, et al. (2012)
European Society for Pediatric Gastroenterology, Hepatology, and Nutrition
guidelines for the diagnosis of coeliac disease. Journal of Pediatrics
Gastroenterology and Nutrition 54: 136160.
Kasarda DD (1994) Toxic cereals in coeliac disease. In: Feighery C and OFarelly C
(eds.) Gastrointestinal Immunology and Gluten-Sensitive Disease, pp. 203220.
Dublin: Oak Tree Press.
Lindfors K, Rauhavirta T, Stenman S, Maeki M, and Kaukinen K (2012) In vitro models
for gluten toxicity: Relevance for celiac disease pathogenesis and development of
novel treatment options. Experimental Biology and Medicine 237: 119125.
Ludvigsson JF, Leffler DA, Bai JC, Biagi F, Fasano A, Green PH, et al. (2012) The Oslo
definitions for coeliac disease and related terms. Gut 62: 4352.
Richey R, Howdle P, Shaw E, and Stokes T (2009) Recognition and assessment of
coeliac disease in children and adults: Summary of NICE guidance. British Medical
Journal 338: b1684.
Sapone A, Bai JC, Ciacci C, Dolinsek J, Green PHR, Hadjivassiliou M, et al. (2012)
Spectrum of gluten-related disorders: Consensus on new nomenclature and
classification. BMC Medicine 10: 13.
Schuppan D, Junker Y, and Barisani D (2009) Celiac disease: From pathogenesis to
novel therapies. Gastroenterology 137: 19121933.
Sollid LM, Qiao S-W, Anderson RP, Gianfrani C, and Koning F (2012) Nomenclature
and listing of celiac disease relevant gluten T-cell epitopes restricted by HLA-DQ
molecules. Immunogenetics 64: 455460.
Wieser H (2007) Chemistry of gluten proteins. Food Microbiology 24: 115119.
Wieser H and Koehler P (2008) The biochemical basis of celiac disease. Cereal
Chemistry 85: 113.
Wieser H, Koehler P, and Konitzer K (2014) Celiac Disease and Gluten:
Multidisciplinary Challenges and Opportunities, 1st edn. London, Waltham, San
Diego: Academic Press.
 The Gluten-Free Diet
V Zevallos, Johannes Gutenberg Mainz, Mainz, Germany
I Herencia, Universidad Politecnica de Madrid, Madrid, Spain
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Human nutrition and the gluten-free diet (GFD).
GFD, celiac disease (CD), and other inflammatory
conditions.
Gluten and nongluten wheat proteins.
Grains with and without gluten.
Legislation of gluten-free products.
Nutritional and technological developments of gluten-free
products.
Learning Objectives
To understand the origin and current situation of a GFD.
To understand the effects of a GFD on health and disease.
To know the current nutritional and technological develop-
ments of GF products.
Introduction
The human diet has changed throughout the years, motivated
mostly by an adaptation to the surrounding environment.
From hunter-gatherers to sedentary populations, the human
diet has evolved, adapted, and changed affecting and influenc-
ing health and disease. Agriculture was one of the main factors
influencing the human diet; for instance, the success of Roman
Empire was arguably supported by agriculture, particularly
wheat production. However, having a diet mostly based in
one staple (wheat) had detrimental effects as indirectly sug-
gested by Aretaeus from Cappadocia, a physician that in AD
200 described a malabsorptive syndrome with chronic diarrh-
ea that was the first account for what is currently known as CD,
an enteropathy affecting mostly the small intestine architecture
in genetically predisposed individuals.
In around 1700, two books described a gastrointestinal
condition characterized by diarrhea (white flux) and the term
sprue from the Dutch spruw was used. At the end of the
nineteenth century, Samuel Gee accurately outlined the typical
features of CD and even highlighted the importance of diet in
the treatment; one of his first dietary intervention was the use
of mussels in a child with suspected CD who relapsed after the
mussels season ended. Haas described Some years ago I
treated a child, aged 3 years, who suffered from a severe case
of anorexia nervosa. She had reached a serious state of deple-
tion and weakness from her self imposed starvation, refusing
all food and regurgitating that fed to her by gavage (tube
feeding). She finally accepted a banana, with the result that
other food was taken in a more or less normal amount within
48 h. There was a complete relapse when the banana was
withheld and food was taken normally only with bananas.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00086-3
According to this observation, Haas described a treatment of
ten celiac patients, eight of which were cured with the banana
diet and two of which died.
The banana diet and other diets based on fruit and vegeta-
bles were the best treatment for celiac patients, but during the
Second World War, those items were unavailable, and
therefore, another dietary treatment was necessary. At that
time, Dicke (Figure 1) and collaborators were the first to
establish a clear link between diet and CD by concluding that
wheat and rye flours caused anorexia, increased fecal output,
and steatorrhea in CD patients, setting up the basis for an
effective treatment. Further studies confirmed their initial
hypothesis by identifying gluten from wheat and related
cereals as the storage proteins triggering CD.
Thus, giving rise to what is currently known as a GFD, this
diet includes naturally occurring gluten-free foods such as fruit,
vegetables, unprocessed meat, fish, and poultry. Products such
as pasta, bread, breakfast cereals, crackers, and snack foods that
are usually manufactured with gluten-containing cereals
(wheat, barley, and rye) are replaced with flour from gluten-
free cereals such as maize, rice, millet, buckwheat, and sor-
ghum. The recent increased prevalence in CD and related
conditions (nonceliac gluten sensitivity (NCGS) and inflam-
matory bowel syndrome (IBD)), the rediscovery of forgotten
highly nutritious staples (quinoa and amaranth), and a recent
social dietary trend have facilitated the expansion of the gluten-
free market.
Current reports indicate that the global gluten-free market
has reached an estimated value of more than $6 billion with a
projected annual growth rate of 10.2% by 2018 (http://www.
marketsandmarkets.com/Market-Reports/gluten-free-products-
market-738.html), suggesting a rapid expansion beyond the
traditional dietary treatment for CD to a global dietary choice
that will compete with traditional food industries. This article
aims to provide an overview about the effects of a GFD on
health and disease, gluten and nongluten proteins and plant
sources, legislation of gluten-free products, and the nutritional
value of a GFD.
Celiac Disease
CD is a gluten-sensitive enteropathy that affects mainly the
villous architecture of the proximal small intestinal in geneti-
cally predisposed individuals (Figure 2). CD is now being
considered a multiorgan autoimmune disease characterized
by an inappropriate immune response triggered by the inges-
tion of amino acid sequences found in seed storage proteins or
prolamins of wheat, barley, and rye. The prevalence of CD
(
1%) was traditionally higher in Europe, where gluten-
containing cereals are frequently cultivated and consumed.
However, increased migration and similar agricultural and
91
92 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet
Figure 1 Professor Willem-Karel Dicke at the Wilhelmina Childrens
Hospital, Utrecht. Reproduced from van Berge-Henegouwen GP and
Mulder CJ (1993) Pioneer in the gluten free diet: Willem-Karel Dicke
19051962, over 50 years of gluten free diet. Gut 34(11): 14731475.
dietary patterns have widen the demographic distribution of
CD with similar prevalence in United States, Australia, South
America, and North Africa where children of Arab-Berber ori-
gin had 5.6% prevalence of antiendomysial antibody. Other
regions (Japan and China) where CD was historically rare are
starting to report new CD cases, which correlates with an
increase in wheat consumption, suggesting that the population
was genetically susceptible but simply did not manifest disease
due to the absence of the environmental factor (dietary gluten)
that is essential to develop CD.
The typical clinical manifestation of CD in adults includes
generalized malabsorption, weight loss, and continuous,
intermittent, or alternated diarrhea with periods of constipa-
tion. In children, classical symptoms such as anorexia,
vomiting, and anemia are observed during the first months of
age, followed by loss of appetite, failure to thrive, behavioral
problems, abdominal distension, small stature, muscle
wasting, and enamel defects during the first years. Moreover,
milder cases characterized by the partial absence of symptoms
or atypical signs are becoming increasingly common, particu-
larly since CD is a complex condition that is associated with
many diseases, not only mainly autoimmune diseases but also
reproductive, neurological, and dermatologic disorders.
The current serological gold standard is IgA endomysial
antibody (EMA), with a specificity that approaches 100% and
a sensitivity of
90%. The titer of EMA correlates with the
degree of mucosal damage. Enzyme tissue transglutaminase
(tTG) was recognized as being the autoantigen for the EMA
allowing the development of an IgA anti-tTG enzyme-linked
immunoassay with less specificity than the IgA EMA test
(9599%) but more sensitivity (9390%). However, histolog-
ical assessment of the small intestine remains the gold standard
in the diagnosis of CD. The celiac lesion predominantly affects
the mucosa of the proximal duodenum with lessening damage
occurring toward the distal part; multiple biopsies using stan-
dard size forceps are needed due to the patchy nature of the
villous changes.
The diagnosis of CD is based on genetic predisposition
(human leukocyte antigen (HLA)-DQ2 or HLA-DQ8), gastro-
intestinal symptoms, and positive serological and histological
abnormalities at duodenal level on a gluten-containing diet.
Once CD has been diagnosed, the main treatment involves
following a strict GFD that consists of eating naturally occur-
ring gluten-free foods such as fruit, vegetables, unprocessed
meat, fish, and poultry. Products such as pasta, bread, breakfast
cereals, crackers, and snack foods that are usually manufac-
tured with gluten-containing cereals (wheat, barley, and rye)
are replaced with flour from gluten-free cereals such as maize,
rice, millet, buckwheat, quinoa, and sorghum. A full list of
gluten-free products is available in Appendix 3. Grains,
Foods, and Ingredients Suiting GFD for CD.
Fortunately, the gluten-free industry has experienced a sig-
nificant development over the last years. However, this rapid
growth has been accompanied by variations in labeling stan-
dards and in the amount of gluten that can be safely tolerated
depending of the country or region, for example, 20 ppm is
considered a safe threshold in Europe. Furthermore, a strict
GFD can only be achieved with difficulties when patients are
traveling and have to eat outside, heavily relying on third
parties (restaurants and shops). In addition, adherence to the
GFD depends mainly on factors such as palatability, availabil-
ity, age, social/peer pressure, and cost. Also, celiac patents that
are extremely sensitive to gluten can have persistent symptoms
despite following a GFD due to consumption of foods contain-
ing hidden sources of gluten such as wheat thickeners, malted
barley with unsafe levels of gluten/ambiguous labels, and other
ingredients such as fermentable starches or some gluten-free
ingredients.
CD requires strict adherence to a GFD for life and hence the
importance to maintain regular contact with a multidisciplin-
ary team formed by the gastroenterologist, GP, dietician, and
nutritionist that can provide ongoing support to the celiac
patients. Failing to maintain a strict GFD could lead to a
persistent stage of inflammation that could be associated to
increased mortality. It is also very important to continue the
development of new products and ingredients to improve the
quality of the GFD.
Other Inflammatory Conditions
There is growing evidence that a GFD has the potential to
become an alternative treatment for gastrointestinal and extra-
gastrointestinal inflammatory conditions such as irritable
bowel syndrome (IBS), NCGS, rheumatoid arthritis (RA), dia-
betes mellitus (DM), and neurological and dermatological
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet 93

Figure 2 Villous architecture of the proximal intestine in a treated (a) and an untreated (b) CD patient, using scanning electron microscopy (left) and
histology (right). Reproduced from Gasbarrini G and Mangiola F (2014). Wheat-related disorders: a broad spectrum of evolving diseases.
United Eur. Gastroenterol. J. 2(4): 254262.

disorders. The pathogenesis of IBS, Crohns disease, and ulcer-
ative colitis remains to be fully elucidated, but genetic predis-
position, microbiome, and environmental/dietary triggers are
the most important factors involved in the onset of disease.
Furthermore, in vivo experiments using HLA-DQ8 mice sensi-
tized with gluten show that an altered barrier function could be
a critical step in facilitating exposure of the mucosal immune
system to dietary antigens that subsequently will trigger an
immune response. Thus, a GFD recommended to patients
with IBD-like symptoms resulted in improved symptoms, sug-
gesting the potential role of a GFD as adjunctive therapy.
Nonceliac gluten/wheat sensitivity (NCGS) is a condition that
significantly improves on a GFD, initially reported in the late
1970s; NCGS can be defined as a reaction to gluten-containing
grains without involvement of allergy or autoimmune
mechanisms. It is associated with gastrointestinal (bloating, diar-
rhea, abdominal pain, and nausea) and predominantly non-
gastrointestinal symptoms (headache, foggy mind, chronic
fatigue, joint and muscle pain, numbness of leg and arms, and
anemia). The prevalence of NCGS is between 1% and 5% and
seems to be more common in females and young/middle-aged
adults although the prevalence in children needs further
investigation. The pathogenic mechanism is still unclear, but
the innate immune response to wheat, particularly part of the
nongluten fraction (amylase trypsin inhibitors, ATIs), seems to
play a significant role in the onset of disease.
RA increases the risk of atherosclerosis and cardiovascular
disease; it has been shown that a gluten-free vegan diet
decreased total cholesterol. It is antiatherogenic and signifi-
cantly reduces disease activity and the levels of antibodies to
b-lactoglobulin. Type I DM is also affected by gluten and
having a GFD has beneficial effects on the preservation of B-
cell function. Wheat consumption also affects neurological
dysfunctions, although the precise mechanism on the nervous
system is still unclear. Thus, a GFD has been shown to improve
and stabilize ataxia by preventing irreversible loss of Purkinje
cells in the cerebellum and improve neuronal sensory action
potential and headaches. Furthermore, there are reports indi-
cating that a GFD has a positive effect on symptom reversal of
multiple sclerosis.
Dietary trends are an important cultural factor often asso-
ciated with health and disease; in recent years. Being on a GFD
has become fashionable in most Western societies. In the
United States, it is considered the most trendy diet with
1.5
million citizens (without any underlying medical condition)
consuming gluten-free products. This proportion is similar in
other Western countries and is predicted to increase even more
in the coming years. The popularity of gluten-free products is
94 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet
partly motivated by the association with healthy eating, weight
loss, and a recent negative perception of wheat as an over-
utilized staple of the Western diet.
Gluten and Nongluten Wheat Proteins
The storage proteins are the main immunogenic components
of wheat, barley, and rye kernels. These proteins are located in
the endosperm of kernels and can be subdivided according to
their solubility in water, salt, alcohol, and alkali into albumins,
globulins, prolamins, and glutelins, respectively. For wheat,
the glutenprotein complex can readily be purified by making
a dough with wheat flour and water and kneading the dough
under running water. The starch and water-soluble proteins
wash out and a cohesive ball of gluten remains. The cohesive
properties of wheat gluten are the reason why only wheat flour
is well suited to bread making. The storage proteins of barley
and rye are similar to those of wheat, but they lack the dough-
forming properties of wheat gluten. For dietary purposes, the
storage proteins of wheat, barley, and rye can be considered as
gluten, being a mixture of both prolamins and glutelins.
Prolamins are alcohol-soluble storage proteins character-
ized by their higher content of proline and glutamine, which
in combination could constitute between 30% and 70% of the
total amino acid composition. The molecular weight of pro-
lamins ranges from 10 to 100 kDa and prolamins are a signif-
icant source of nitrogen, sulfur, and carbon for the developing
embryo and they are known as gliadins in wheat, hordeins in
barley, and secalins in rye.
According to their decreasing electrophoretic mobility, gli-
adins can be divided into a, b, g, and o gliadins. Based on their
N-terminal amino acid sequences, gliadins can be categorized
into a, g, and o. There is another classification based on the
amount of sulfur, o gliadins are sulfur-poor and a and g
gliadins are sulfur-rich. The first prolamin to be fully
sequenced was A-gliadin (an a-type) using direct protein
sequencing; the remaining amino acids were sequenced using
cloned DNA, facilitating the development of synthetic pep-
tides. Other studies using different wheat, barley, and rye cul-
tivars found that purified prolamins from rye (g and o
secalins) and from barley (o hordeins and C-hordeins) could
be identified using chromatographic methods. Prolamins are
also present in other plants but at a different proportion
and with different amino acid sequences than in wheat, barley,
and rye.
The glutelin protein of wheat is termed glutenin. It consists
of alcohol-soluble subunits that are cross-linked by interchain
disulfide bonds to form alcohol-insoluble polymeric glutenin.
These subunits have been classified according to their electro-
phoretic mobility into high-molecular-weight (HMW) and
low-molecular-weight groups. The HMW glutenin subunits
make a major contribution to dough formation. Specific sub-
units are identified according to the relevant genes, such as
1Dy10, 1Dx5, 1Dx4, and 1Dy9. Rye and barley have equiva-
lent glutelin fractions (HMW secalins and D-hordeins, respec-
tively) but they are less suitable for dough formation.
Immunogenic gluten (prolamins and glutelins) peptides
trigger an adaptive immune response in CD. These peptides
reach the lamina propria via a combination of defective tight
junctions, a paracellular pathway, epithelial transcytosis, and
direct sampling by dendritic cells. Internalized gluten peptides
are then deamidated by the enzyme tTG II, increasing nega-
tively charged peptides that can be accommodated in binding
pockets of the heterodimeric HLA class II alleles encoding for
HLA-DQ2 and HLA-DQ8. This increase in antigenicity acceler-
ates the secretion of interferon-g (IFN-g) by gluten-specific
CD4 T-helper cells, contributing to the expansion of
cytotoxic T cells and fibroblasts and the release of metallopro-
teases to degrade the extracellular matrix of enterocytes at the
lamina propria and also to start secretion of anti-tTG by gluten-
specific plasma cells into the lamina propria and then to the
bloodstream.
The other major grouping of wheat-grain proteins is the
water-/salt-soluble proteins referred as nongluten wheat
proteins, the most relevant in terms of innate inflammation
being the ATIs. Some of these proteins play a significant role as
a plant defense mechanism against pests and during the met-
abolic process of seed development. Wheat ATIs were identi-
fied as contaminating proteins within pepsintrypsin gliadin
digests; they represent between 2% and 4% of total grain
protein compared to 7080% for gluten. ATIs are resistant to
proteolytic digestion due to their compact secondary structure
that includes five intrachain disulfide bonds. ATI subfamilies
can be monomeric or form noncovalently linked dimers or
tetramers. There are up to 17 ATI species of around 150-amino
acid length with a variant primary structure and conserved
secondary structure.
ATIs engage the toll-like receptor 4 (TLR4)-MD2-CD14
complex and lead to upregulation of maturation markers and
release of proinflammatory cytokines in myeloid cells of celiac
patients. In vivo feeding experiments in mice showed that ATIs
increased intestinal and systemic release of cytokines and che-
mokines like IL-8, TNF-a, and CCL-2 within 212 h, and
ex vivo celiac biopsy studies demonstrated an adjuvant effect
of added ATIs for the gluten-specific T-cell response. Therefore,
ATIs appear to be the sought triggers of innate immunity in CD
and NCGS.
Gluten-Containing and Gluten-Free Plants
Gluten-containing cereals are part of the Gramineae family,
also known as Poaceae due to its association with the type
genus Poa. This family includes 12 subfamilies, in which two
are more relevant to CD: Pooideae that includes wheat, rye,
and barley, which are considered toxic for celiac patients, and
Panicoideae that includes maize, millet, and sorghum, which
are considered suitable for the GFD. The food product corn
gluten is a misnomer; it is the protein washed from maize
(corn) grain during wet milling. It is not related to wheat
gluten.
The taxonomic plant classification proposed by Kasarda
provided a simple guidance to separate safe from unsafe grains.
Monocotyledonous plants, members of the Gramineae
family, and the Triticeae tribe (wheat, barley, and rye) were
considered celiac-toxic, while dicotyledonous plants such as
Chenopodiaceae and Amaranthaceae (quinoa and amaranth)
and those from other tribes (maize, rice, and tef) are consid-
ered safe for celiac patients.
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet
Early studies on oats, probably contaminated with gluten,
indicated that oats were unsuitable, but recent feeding studies
comparing a GFD with and without oats concluded that mod-
erate amount of oats (5070 g per day) can be safely added to
the GFD of celiac patient. Another study assessing the long-
term effects of large amount of oats (up to 100 g per day)
found that histology, serology, and well-being were constant
throughout the study. However, there have also been reports
that activation of CD with morphological deterioration
occurred after oats challenge. They described T-cell lines reac-
tive to avenin (the storage protein of oats) from duodenal
mucosa of five patients consuming oats as part of their GFD
that were positively stimulated by avenin peptides rich in
proline and glutamine residues. Other reports indicated with-
drawals from some large feeding studies, which added to the
T-cell data, suggest that for a minor proportion of the celiac
population, oats are unsuitable.
Furthermore, gluten-containing plants (wheat, rye, barley,
and their earlier ancestors) rich in ATIs have a strong innate
stimulatory capacity that contrasts with a rather minimal
innate stimulatory effect of ATIs from gluten-free plants.
Thus, gluten-free plants are also ATI-free. The average amount
of wheat flour consumed per adult/day is
250 g, indicating
that around 1 g of ATIs could be enough to trigger a moderate
innate immune response in the gut of people affected by CD or
NCGS, also having an effect on nongastrointestinal organs.
The morphological effect of ATIs is discrete and characterized
by an increase in inflammatory cells in accordance to histolog-
ical changes observed in NCGS patients. Furthermore,
the effect is dose-dependent, and therefore, a reduction
(8090%) of daily ATI consumption may be sufficient to elim-
inate the stimulatory effect, and therefore, a strict GFD does
not need to be maintained. Over the last decades, an increased
prevalence of allergies and autoimmune diseases has been
observed, particularly in developing countries. This can be
partly explained by a parallel increase in wheat (ATIs) con-
sumption since traditional (gluten-free) staples were replaced
by more affordable alternatives (modern wheat).
Legislation of Gluten-Free Products
The denomination gluten-free theoretically implies without
any amount of gluten; however, having such a diet will be
impractical and very difficult to implement since gluten is a
ubiquitous ingredient and therefore will be almost inevitably
present in trace amounts within any diet. Furthermore, com-
pliance to a GFD is mainly related to palatability and the
availability of gluten-free products. This difficulty was partly
addressed by the addition of wheat starch flour to the gluten-
free list, particularly in Scandinavia and the United Kingdom
where patients consumed traces up to 200 ppm without clini-
cal evidence of exacerbation of the disease. However, there is
an individual level of tolerance to gluten and even small
amounts of gluten can be of relevance for a group of celiac
patients.
The other alternative to increase palatability and reduce the
risk of nutritional imbalance is to include alternative naturally
gluten-free ingredients. However, there is a risk of gluten con-
tamination that could occur from harvest to packaging, for
example, samples of millet, sorghum, and soy heavily
95
contaminated with gluten (up to 3000 mg kg1). Although
these products were not labeled as gluten-free, they still repre-
sent a risk for celiac patients, particularly since clinical studies
have suggested that 100 mg kg1 of gliadin or more can trigger
an inappropriate immune response leading to histological
damage of the intestinal mucosa in diagnosed patients.
Similarly, it was found that the daily intake of gliadin at
414 mg did not produce the same effects for most celiac
patients.
The codex standard for gluten-free foods (Codex Stand 118,
1981) regulates the level of gluten in gluten-free foods and is
often reviewed by a special FAO/WHO committee. The last
review of Codex Alimentarius Commission (2008) indicates
that the maximum gluten level for gluten-free products is
20 mg kg1. In 2012, new legislation based on these guidelines
was issued by the European Commission, and the 20 mg kg1
threshold was applied to all GF products, that is, naturally GF
(maize, rice, and uncontaminated oats) and derived from
wheat starch. Other organizations such as the Food and Drug
Administration have adopted similar a similar threshold level
(20 ppm), and the Food Standards Australia New Zealand has
chosen a more strict approach considering no detectable
amount of gluten or as low as 35 ppm (www.standards.gov.
au; www.coeliac.org.au).
Nutritional Value of the GFD
Restrictive diets such as a GFD tend to have an inherent risk of
nutritional deficiency. Previous studies mostly agreed that a
GFD is nutritionally inadequate, but other studies did not
reach the same conclusion (http://www.food.gov.uk/science/
research/allergy-research/t07053), and in general, information
about the nutrition adequacy of a GFD is still rather limited.
There is evidence that adults following a GFD have
increased levels of cholesterol, altered homocysteine levels,
and a potentially high risk of developing cardiovascular dis-
ease. Furthermore, another study showed that gluten-free
products, particularly breads and pasta, have low protein,
high saturated fat, sodium, and cholesterol compared to
gluten-containing products, reflecting the nutritional conse-
quences (low protein/high fat) particularly in females.
This nutritional imbalance can be due to the ingredients
used. For instance, wheat starch requires an industrial process
that inevitably alters the macro- and micronutrient composi-
tion of wheat, a source of protein, iron, and vitamins (thiamin,
riboflavin, folate, and niacin). In addition, replacing tradi-
tional gluten-free products for alternative naturally gluten-free
products such as Andean grains (quinoa and amaranth) can
increase variety, palatability, and most importantly significant
amounts of nutrients such as protein, fiber, iron, calcium,
vitamins, and polyunsaturated fatty acids.
Baking gluten-free products remains a challenge mainly
because the main ingredient, gluten, cannot be easily replaced.
Most of the gluten-free flours lack the viscoelastic properties of
wheat dough when kneaded in water and usually require more
water to form a dough, which is difficult to handle. It is often
referred to as a batter. The main problem arises when the
mixture of water, yeasts, gluten-free flours, and starches forms
few coherent structures because the gases from fermentation
are not entrapped, directly affecting quality parameters, having
96 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet
reduced volume and crumbly structure, with poor texture,
mouthfeel, and flavor.
However, the gluten-free baking industry through food
technologists and nutritionists maintains a continuous process
of quality development, particularly by the addition of novel
ingredients and new technologies. Thus, the main goal is to
create gluten-free products with similar quality standards to the
gluten-containing counterparts that will ultimately help to
improve the quality of life of people affected by wheat-related
conditions.
Conclusion and Future Developments
Patients affected by wheat and related cereals benefit from a
GFD. There is clear evidence that gluten proteins exacerbate
celiac disease. However, non-gluten proteins as activators of
innate immunity might also play a significant role in CD and
other inflammatory conditions. Health care professionals sup-
port patients on a GFD by improving their psychological well-
being, reducing social restrictions, developing new products to
meet nutritional targets, and ultimately improving their quality
of life. The increased number of people benefited from a GFD
as well as a recent social trend indicates that the gluten-free
market is projected to grow significantly over the coming years.
Exercises for Revision
Describe the most common symptoms of CD for children
and for adults.
What are the screening tests and the gold standard tests to
diagnose CD?
Outline the taxonomic groupings of grain species that are
celiac-toxic or not. Refer to the encyclopedias article on the
taxonomy of grain-bearing plants.
Summarize the development of legislation to assist con-
sumers in choosing gluten-free foods, also considering the
means of communication, mainly via product labeling.
Exercises for Readers to Explore the Topic Further
Investigate the recorded evidence for oats being harmful or
not in CD. Refer to recommendations in Appendix 3, and
examine articles such as the following:
- Kemppainen, T.A. et al., 2008. Unkilned and large
amounts of oats in the coeliac disease diet: a randomized,
controlled study. Scandinavian Journal of Gastroenterology,
43(9), pp. 10941101.
- Lundin, K.E.A. et al., 2003. Oats induced villous atrophy
in coeliac disease. Gut, 52 (11), pp. 16491652.
In recent years, a GFD has become fashionable in many
societies. Investigate the existence of this fashion in your
region. Determine the extent to which this trend, if appar-
ent, has a cultural-fashion basis or if it may be associated
with valid health and disease reasons.
In parallel, investigate the availability of gluten-free foods
in your region, considering over-the-counter goods and
packaged foods in the local shops.
Investigate the methods used to determine the amount of
gluten in food products and ingredients. Start with the
encyclopedias articles on the standardization of methods
for analyzing grains and Appendix 3, which lists testing
methods.
See also: Appendix 3: Grains, Foods, and Ingredients Suiting
Gluten-Free Diets for Celiac Disease; Food Grains and the
Consumer: Consumer Trends in Grain Consumption; Grains and
Health: Misinformation and Misconceptions; Labelling of Grain-Based
Foods; Food Grains: Intolerance, Allergy and Diseases: Celiac
Disease; Cereal Allergens; Proteins: The Protein Chemistry of Cereal
Grains; The Basics: Taxonomic Classification of Grain Species; The
Cereal Grains: Wheat: An Overview of the Grain That Provides Our
Daily Bread; Wheat-Based Foods: Cereals: Overview of Uses:
Accent on Wheat Grain.
Further Reading
Catassi C, et al. (1993) Dose dependent effects of protracted ingestion of small amounts
of gliadin in coeliac disease children: A clinical and jejunal morphometric study. Gut
34(11): 15151519.
Catassi C, et al. (1999) Why is coeliac disease endemic in the people of the Sahara?
Lancet 354(9179): 647648.
Catassi C, et al. (2013) Non-celiac gluten sensitivity: the new frontier of gluten related
disorders. Nutrients 5(10): 38393853.
Ciacci C, et al. (1999) Low plasma cholesterol: A correlate of nondiagnosed celiac disease
in adults with hypochromic anemia. Am. J. Gastroenterol. 94(7): 18881891.
Dicke WK, Weijers HA, and Van De Kamer JH (1953) Coeliac disease. II. The presence
in wheat of a factor having a deleterious effect in cases of coeliac disease. Acta
Paediatr. 42(1): 3442.
Dieterich W, et al. (1997) Identification of tissue transglutaminase as the autoantigen of
celiac disease. Nat. Med. 3(7): 797801.
Dupont FM, et al. (2011) Deciphering the complexities of the wheat flour proteome
using quantitative two-dimensional electrophoresis, three proteases and tandem
mass spectrometry. Proc. Natl. Acad. Sci. U. S. A. 9: 10.
Ejderhamn J, Veress B, and Strandvik B (1988) The long-term effect of continual
ingestion of wheat starch-containing gluten-free products in coeliac patients.
In: Kumar PJ (ed.) Coeliac Disease: One Hundred Years, pp. 294297. Leeds: Leeds
University Press.
Gasbarrini G and Mangiola F (2014) Wheat-related disorders: a broad spectrum of
evolving diseases. United Eur. Gastroenterol. J. 2(4): 254262.
Haas SV (1924) The value of banana in the treatment of celiac disease. Arch. Pediatr.
Adolesc. Med. 28(4): 421.
Hadjivassiliou M, et al. (2003) Dietary treatment of gluten ataxia. J. Neurol. Neurosurg.
Psychiatry 74(9): 12211224.
Hafstrom I, et al. (2001) A vegan diet free of gluten improves the signs and symptoms of
rheumatoid arthritis: The effects on arthritis correlate with a reduction in antibodies
to food antigens. Rheumatology 40(10): 11751179.
Hager AS, et al. (2012) Investigation of product quality, sensory profile and
ultrastructure of breads made from a range of commercial gluten-free
flours compared to their wheat counterparts. Eur. Food Res. Technol. 235(2):
333344.
Herfarth HH, et al. (2014) Prevalence of a gluten-free diet and improvement of clinical
symptoms in patients with inflammatory bowel diseases. Inflamm. Bowel Dis. 14.
Junker Y, et al. (2012) Wheat amylase trypsin inhibitors drive intestinal inflammation via
activation of toll-like receptor 4. J. Exp. Med. 209(13): 23952408.
Kasarda DD (1994) Toxic cereal grains in coeliac disease. In: Feighery C and
OFarrelly C (eds.) Proceedings of the 6th International Symposium on Coeliac
Disease, pp. 203220. Dublin: Oak Tree Press.
Kasarda DD, et al. (1984) Nucleic acid (cDNA) and amino acid sequences of alpha-type
gliadins from wheat (Triticum aestivum). Proc. Natl. Acad. Sci. U. S. A. 81(15):
47124716.
Kemppainen TA, et al. (2008) Unkilned and large amounts of oats in the coeliac disease
diet: a randomized, controlled study. Scand. J. Gastroenterol. 43(9): 10941101.
 FOOD GRAINS: INTOLERANCE, ALLERGY AND DISEASES | The Gluten-Free Diet
Lee AR, et al. (2009) The effect of substituting alternative grains in the diet on the
nutritional profile of the gluten-free diet. J. Hum. Nutr. Diet. 22(4): 359363.
Lo W, et al. (2003) Changing presentation of adult celiac disease. Dig. Dis. Sci. 48(2):
395398.
Lundin KEA, et al. (2003) Oats induced villous atrophy in coeliac disease. Gut 52(11):
16491652.
Miranda J, et al. (2014) Nutritional differences between a gluten-free diet and a diet
containing equivalent products with gluten. Plant Foods Hum. Nutr. 69(2):
182187.
Molberg O, et al. (1998) Tissue transglutaminase selectively modifies gliadin peptides
that are recognized by gut-derived T cells in celiac disease. Nat. Med. 4(6): 713717.
Osborne TB (1907) The Protein of the Wheat Kernel. Washington, DC: Carnegie Institute
of Washington, 84.
Pastore MR, et al. (2003) Six months of gluten-free diet do not influence autoantibody
titers, but improve insulin secretion in subjects at high risk for type 1 diabetes.
J. Clin. Endocrinol. Metabol. 88(1): 162165.
Pengiran Tengah CDSNA, et al. (2004) Multiple sclerosis and occult gluten sensitivity.
Neurology 62(12): 23262327.
97
Peraaho M, et al. (2003) Wheat-starch-based gluten-free products in the treatment of
newly detected coeliac disease: prospective and randomized study. Aliment.
Pharmacol. Ther. 17(4): 587594.
Schuppan D, Junker Y, and Barisani D (2009) Celiac disease: from pathogenesis to
novel therapies. Gastroenterology 137(6): 19121933.
Shewry PR and Tatham AS (1990) The prolamin storage proteins of cereal seeds:
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513519.
Strsrud S, et al. (2003) Adult coeliac patients do tolerate large amounts of oats. Eur. J.
Clin. Nutr. 57(1): 163169.
Zanini B, et al. (2013) Impact of gluten-free diet on cardiovascular risk factors. A
retrospective analysis in a large cohort of coeliac patients. Dig. Liver Dis. 45(10):
810815.
Zevallos VF, Herencia LI, and Ciclitira PJ (2014) Gastrointestinal effects of eating
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109(2): 270278.
 Proteins
Contents
The Protein Chemistry of Cereal Grains
The Protein Chemistry of Dicotyledonous Grains
Protein Synthesis and Deposition
Nitrogen Metabolism
Nitrogen in Grain Production Systems
The Enzymes Associated with Food Grains and Their Functions in Processing
Proteomic Analysis Relevant to Cereal Grains
Thiolomics of the Gluten Protein Network of Wheat Dough
The Protein Chemistry of Cereal Grains
F Bekes, FBFD PTY LTD, Beecroft, NSW, Australia
CW Wrigley, The University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The importance of dietary protein from cereal grains rice,
wheat, maize, and others.
The purification of wheat gluten from dough as an exper-
iment in the kitchen and as a commercial product.
The simple structure of proteins a linear polymer of
amino acids produces the diversity of properties
of enzymes, hormones, hair and horn, silk and sinew,
and gluten together with the many other grain storage
proteins.
Folding and cross-linking of the amino acid polymer gives
three-dimensional structure that is needed for many pro-
tein functions, such as enzymatic activity.
Osbornes solubility-based protein classes: water-soluble
albumins, salt-soluble globulins, prolamins extractable
with 70% aqueous ethanol, and the residual glutelins.
Protein quantity is determined by various methods, espe-
cially by near-infrared spectroscopy, based on total nitrogen
content.
Protein quality relates to its nutritional value and its con-
tribution to functional uses.
The specific proteins are described for wheat, rye, triticale,
barley, rice, maize, and sorghum.
Learning Objective
To achieve an understanding of proteins as critical compo-
nents of the grains of cereal species and their function for
the plants and for our dietary needs.
98 Encyclopedia of Food Grains, Second Edition
Introduction
Worldwide, plant proteins contribute more than half of our
dietary supply of protein. Over 200 million tonnes of protein
are harvested each year in the form of cereal grains, based on FAO
statistics (http://faostat.fao.org). This protein harvest includes
that derived from the three major cereal grains, wheat, rice, and
maize, and also less widely grown species: barley, rye, triticale,
and sorghum. In some developing countries, a single cereal
species may be the major or sole source of protein in the diet.
In many countries, cereal proteins enter the diet in various
forms, including wheaten breads and pastries, noodles, fried
rice dishes, rye crispbreads, cookies, biscuits, pearl barley,
pasta, corn tortillas, breakfast cereal, oatmeal porridge,
couscous, and many more foods. Most obviously, cereal grain
proteins are eaten directly in these many processed forms. The
second route is indirect, in the form of animal products, such
as meat, milk, and eggs, following the feeding of cereal grains
to animals.
Whereas the protein part of the grain is important nutri-
tionally, it also serves functional purposes, relevant to the
processing of the grain. Of these, wheat and rye are unique
because of the dough-forming properties of their storage pro-
teins, known as gluten when formed into dough.
The Discovery of Protein Structure
Cereal Proteins in Prehistory
Historically, the proteins of the cereal grains have played a
significant part in the civilization of mankind, through their
role in providing a reliable source of amino acids, the essential
http://dx.doi.org/10.1016/B978-0-12-394437-5.00101-7
 PROTEINS | The Protein Chemistry of Cereal Grains 99

dietary compounds that are the building blocks of proteins

in the body. Grains as a source of dietary proteins could be
obtained via agriculture, permitting the establishment of a
permanent dwelling place, providing an important change
from the nomadic existence of the hunter-gatherer way of
life. The establishment of permanent towns led, in turn, to
specialization of occupation and to opportunities for mankind
to pursue cultural pursuits and scientific inquiry. However, it is
only in recent centuries that the chemistry of proteins has been
elucidated.

Early Concepts
The word protein was proposed by the French scientist Berze-
lius, writing to the German scientist Mulder, with the appro-
priate meaning of primary substance, as recently as 1838, long
after the acceptance of the terms for specific types of proteins,
such as gluten, zymom, and fibrin. Early concepts of protein
chemistry imagined it to have the complex formula
C20H31N5O6, meaning that 20 molecules of carbon (C) were
compounded with 31 molecules of hydrogen, five molecules
of nitrogen, and six molecules of oxygen. The dogma of those
early days also maintained (incorrectly) that this complex was
absorbed (unchanged) directly by the intestine. It is now
known that dietary proteins are hydrolyzed to the component
amino acids for absorption into the bloodstream.

Wheat Gluten: The First Purified Protein

One of the first proteins to be obtained in reasonably pure
form was gluten, because it could be prepared so readily (by
the washing of dough made from wheat flour). This important
experiment, first reported by the Italian scientist Beccari in
1729, can be simply performed in the home kitchen. To do
so, a dough is mixed by adding water, a little at a time, to wheat
flour until a cohesive mass is formed. This dough is kneaded
between the fingers under a gentle stream of water from the
tap. If a glass is placed underneath, a milky-white stream of
starch can be caught in the glass, while the dough in the fingers
becomes smaller and tougher. The gluten ball held in the
fingers is mainly protein, separated very simply from the starch
that settles at the bottom of the glass.
Strictly speaking, gluten is a proteinlipidcarbohydrate
complex formed as a result of specific covalent and non-
covalent interactions from flour components during dough
making as the components are hydrated and energy from
mechanical input from the mixing process is provided.
This gluten-washing procedure has since become very big
business internationally, because the gluten thus isolated is
needed for bolstering the protein content of a wide range of
food products bread, especially, and also breakfast cereals,
meats, cheeses, snack foods, and texturized products or con-
sumed directly as seitan. Nonfood applications include pet
and aquaculture feeds, biodegradable plastics, films, coatings,
adhesives, inks, cosmetics, and pharmaceuticals.
Late in the nineteenth century, the American scientist
Thomas Osborne studied the proteins of many edible grains,
using a succession of solvents to extract diverse classes of pro-
teins from the crushed grain. As a result, he proposed the
generic names of albumin (extracted with water), globulin exist in the grain in monomeric or polymeric form. In the case
(extracted subsequently by salt solution), prolamin (extracted
Figure 1 The cover of one of Osbornes early publications (1912) on
the proteins of cereal grains.
by 70% aqueous ethanol), and glutelin (extracted finally by a
dilute acid). Subsequently, the prolamins of wheat have been
understood to be single polypeptides, whereas wheat glutelins
(known as glutenins) have been shown to be disulfide-linked
polymers built up of glutenin subunits. The cover of one of
Osbornes famous publications appears in Figure 1.
In many cereal grains, the metabolic proteins are albumins
and globulins, many of these being the residual enzymes that
have functioned in synthesizing the components of the matur-
ing grain, plus others that are present in preparation for germi-
nation. On the other hand, the cereal storage proteins are often
of the prolamin and glutelin classes; their water insolubility
offers the advantage that they will be retained when the grain
becomes waterlogged in the germination process.
Osbornes names for these major protein classes have since
continued in use, but their definitions have been refined, and it
has become clear that any one of these protein classes contains
a large number of distinct but related proteins. The modern
classification of cereal proteins is based on their genetics, also
based on the chromosomal location of their coding genes. The
term prolamin in this nomenclature system covers all the
polypeptides soluble in ethanol, independently whether they
of wheat (Figure 2), this prolamin group is subdivided into three
100 PROTEINS | The Protein Chemistry of Cereal Grains
Wheat proteins
Soluble proteins
Monomeric gliadin
Albumins Globulins gliadins
gliadins gliadins
S-poor
prolamins
HMW prolamins
HMW
glutenin subunits
LMW
glutenin subunits
S-rich prolamins
Figure 2 Classification of the proteins of the wheat grain, showing the electrophoretic patterns for some of them.
groups as sulfur-rich, sulfur-poor, and high-molecular-weight
(HMW) prolamins, representing the a-, b-, and g-gliadins
plus the low-molecular-weight (LMW) glutenin subunits, the
o-gliadins, and finally the HMW glutenin subunits, respectively,
in the old classification system.
So What Is a Protein?
A protein molecule is simply a linear polymer of amino acids.
The structure of protein molecules also depends on the folding
and cross-linking of the polymers, giving a three-dimensional
structure that is especially important for functions such as
enzymatic activity.
It is no wonder that these early protein chemists wrestled
with reaching this understanding, when we now see that this
relatively simple chemical structure can produce the great
diversity of properties that are found in proteins, such as
enzymes and epidermis, hair and horn, feathers and flagella,
silk and sinew, and even spiders web.
The diversity depends on the selection and sequence of the
amino acids plus the length of the protein (polypeptide) chain.
Further diversity is provided by cross-linking between polypep-
tide chains and from linking to accompanying molecules
(lipids, carbohydrates, or metal ions). The resulting protein
molecules are large compared with many biological com-
pounds, with molecular weights often in the range of
10 000100 000 Da (1 Da being equivalent to the mass of a
hydrogen atom). Some proteins are even larger, the largest
glutenin proteins of wheat gluten having sizes extending up
to tens of millions of Daltons.
Gluten proteins
Polymeric glutenin
LMW HMW
glutenin subunits
glutenin subunits
gliadins
S-rich HMW
prolamins prolamins
S-poor prolamins
gliadin
gliadin
gliadin
gliadin
There are about 20 different amino acids, each of them
having the combination of an acidic group (
COOH) and a
basic group (
NH2) and each having a distinct side chain that
provides the functional differences between them all. The
amino acids are joined together by peptide bonds to form a
polypeptide chain, by the following formula (for two amino
acids, R1 and R2):
R 1
COOH H2 N
R 2 ! R 1
CO
NH
R2 H2 O
The sequence and specificity of the amino acids are spelled
out exactly in the genetic code of the genes, thereby ensuring
that the protein will be made in the same manner each time
in the specific species and tissue. For this reason, the determi-
nation of grain protein composition is a valuable means of
establishing the identity of a variety within a grain species.
Many Different Cereal Grain Proteins
In the past century, we have come a long way from Osbornes
concept of four classes of proteins to realize that within each of
his classes, there are a large number of proteins differing in
charge and size; this great diversity and heterogeneity can
readily be shown by fractionation methods such as gel electro-
phoresis. Each of the horizontal lines (bands) in the electro-
phoretic patterns in Figure 3 represents a separate protein
component (more than one in many cases). The combination
of gel electrophoresis and gel isoelectric focussing as used in
proteomics, provides a two-dimensional display of spots
(Figure 4), each being a distinct protein (polypeptide, mean-
ing polymer of amino acids).
 PROTEINS | The Protein Chemistry of Cereal Grains 101

Figure 3 Lab-on-a-chip analysis of the composition of prolamin proteins extracted from grains of wheat varieties (L to R: Halberd, Janz, Frame, and
Yitpi) and of barley varieties (Excalibur, Sloop, Schooner, Flagship, Gairdner, and Baudin). Each protein extract was applied to a small lab chip for
microfluidic capillary electrophoresis. Each analysis took about 50 s. The elution patterns were manipulated to simulate gel electrophoresis patterns.

3 pH 10
kDa
130

10
Figure 4 Proteome map of the polypeptides (reduced proteins)
extracted from mature wheat grain (cv Rosella). Fractionation involved
isoelectric focussing between pH 3 (left) and pH 10 followed by SDS gel
electrophoresis in a gradient from 8% (top) to 18% polyacrylamide.
After in-gel digestion with trypsin, 315 protein spots (as numbered) were
excised and identified by MALDI-TOF and LCMS/MS. Adapted from
Mak, Y., Willows, R. D., Roberts, T. H., Wrigley, C. W., Peter Sharp, P.,
and Copeland, L., 2009. Germination of wheat: A functional proteomics
analysis of the embryo. Cereal Chemistry 86 (3), 281289.
The diversity of grain protein composition is such that
differences in composition can readily be demonstrated
between closely related varieties of a grain species. This is
evident in Figure 3 where each column of bands represents
Crude protein Crude fat Crude fiber
Crude ash Starch Cellulose
Figure 5 Chemical composition typical of a whole cereal grain.
the prolamin proteins extracted from grain samples of specific
varieties of wheat and of barley. Each of the two sets of
varieties has a distinct pattern of bands, while all the wheat
patterns differ from all the barley patterns. This type of anal-
ysis is thus useful for the identification of varieties in the
practical situation of grain delivery when identity must be
established to define quality type and royalty payment for
distinct varieties.
102 PROTEINS | The Protein Chemistry of Cereal Grains

Why Is Protein So Important in Grains? and the fiber components (Figure 5). The protein content of
cereal grains ranges from 8% to 15% of the grain weight (Table 1).
The protein component of the grain is critical to the grains role Even within a grain species, the actual protein concentration may
as a seed the beginning of a new plant. The grain protein has cover a range of values, depending on the genetic potential of a
the natural purpose to serve as a reserve of amino acids for the specific variety of that grain species, on the nutritional status of
embryo (germ) to draw upon during germination, while the the plant, and on climatic and other environmental factors.
roots and shoots are growing until the leaves can support Most of the protein is in the grains endosperm (the part
growth and development via photosynthesis. that becomes white flour after milling the cereal grain), the
The protein component of the grain is important nutrition- endosperm being the largest part of the grain by mass. How-
ally as a source of amino acids, especially the essential amino ever, the proportion of protein may be higher in some of the
acids that we cannot synthesize ourselves. The amino acids that non-endosperm tissues of the grain, especially in the germ (the
are essential include lysine, methionine, leucine, isoleucine, embryo, which will become the new plant after germination).
and several more. The barley grain is taken as an example of this in Figure 6.
On the other hand, grain protein is important for the grain Figure 7 shows the chemical composition of wheat grain, with
processing industry, because it is often the most important part of respect to the distribution of protein (and other components)
the grain in determining processing quality and nutritional value. in its various anatomical parts.
In grain composition, protein is next in abundance after starch The importance of protein content is seen in the valuation
of grain shipments. The single figure of the protein content of
a grain sample is an important specification in the marketing
Table 1 Approximate protein content of the more common cereal of most cereal grains. In general, grain of higher protein con-
species, as percentages of the as is weight of whole grain, except for tent commands a higher value. For wheat, a higher protein
rice (milled grain)
content indicates better baking quality, and a higher protein
Cereal species Protein content content for feed grains provides a richer supply of essential
amino acids (irrespective of the proportion of essential amino
Barley 912 acids in their storage protein).
Maize (corn) 812 However, malting barley is an exception to the rule that
Oats (as groats) 1215 high protein content is desirable, because of the need to have a
Rice (milled) 811 high starch content to provide a correspondingly high extrac-
Rye 1215
tion of fermentable sugars in the malting process. Nevertheless,
Wheata 916
even for malting barley, there is a minimum level of protein
a
Durum wheat, a distinct species from common wheat, generally requires a higher (8%), to ensure that there is an adequate supply of the hydro-
protein range, say, 1216%. lytic enzymes needed throughout the malting process.

Crude protein

Crude fat

Crude fiber

Crude ash

Nitrogen-free extract

Cellulose

Crude fiber Crude fat Crude protein

20 5 13.5
4 13.0
15
12.5
3
10 12.0
2
11.5
5 1 11.0
0 0 10.5
Whole Pearled Hulls Whole Pearled Hulls Whole Pearled Hulls
Figure 6 Chemical composition of the barley grain; distribution between the different parts.
 PROTEINS | The Protein Chemistry of Cereal Grains 103

12% Protein
2% Lipids
60% Starch
2% Red. sugars
6% Pentosans
% of grain 2% Cellulose
2% Ash
0 50 100
Embryo Scutellum Pericarp Aleurone Endosperm
Protein
Lipids
Starch
Red. sugars
Pentosans
Cellulose
Ash

Figure 7 Chemical composition of the wheat grain; distribution between anatomical parts. Red. Sugars means reducing sugars, such as glucose.

Table 2 Methods used to determine the protein content of cereal elements and determination of the proportion of the element
grain, especially wheat nitrogen. This figure is multiplied by 6.25 to obtain the protein
content or by a factor of 5.7 in the case of wheat. This proce-
Process Material assayed Assay method dure assumes that virtually all the nitrogen in the grain is in the
form of protein. A lower multiplication factor is used for wheat
Grain NIR
Mill Flour/meal NIR
because its protein contains a high proportion of the amino
# acid glutamine, giving its protein unusually high nitrogen
Wash (for wheat Gluten Weight (wet or dry) content (Figure 8). The total nitrogen content is used as an
only) absolute value in establishing correlations for the routine
# method of NIR spectroscopy to determine the protein content
Extract Protein in solution Biuret or dye-binding of whole grain or of milled products (Table 2).
reaction
#
Hydrolyze Amino acids Gas or liquid Protein Quality
chromatography
# The single figure of protein content, however, does not take
Alkaline digestion Amide N Titrate ammonia released into account the quality of the protein. This quality is indicated
#
for nutritional purposes by analysis of the levels of essential
Acid digestion Ammonia or Kjeldahl or Dumas method
amino acids after hydrolysis of the peptide bonds. For wheat,
nitrogen
however, the quality of the protein also relates to the dough
strength provided by the particular combination of gluten pro-
teins present, determined partly by the genotype (the variety)
and also by the growth conditions.
Analysis of Protein Content For the animal feed industry, the amino acid composition is
especially important, due to the need to ensure an adequate
Today, the routine determination of protein content of all supply of essential amino acids. The protein of cereal grains is a
grain species generally involves the use of near-infrared (NIR) good source of most of the essential amino acids, with the
spectroscopy, applied to whole grain, milled grain, or flour, exception of lysine and possibly tryptophan, which are lower
based on correlations to total nitrogen analyses (Table 2). In in the cereal grains than in most animal proteins. The amino
the case of wheat only, protein content can also be determined acid composition for wheat is shown in Figure 8, with respect
by washing out the gluten protein and weighing its wet mass. to the major classes of wheat proteins. The amino acid com-
Even though gluten protein is less than 80% of the total grain positions of whole grains of several cereals are compared in
protein, this approach can be used if appropriate correlations Table 3.
are established between the level of washed gluten and true
grain protein content.
Further possibilities include colorimetric methods (e.g., the Comparison of the Proteins of Cereal Species
biuret reaction) or the amide distillation procedure (Table 2).
The standard (true) method for protein determination The various cereal species differ in the composition of their
involves the complete digestion of the grain or flour to its grain proteins. Just as the various cereal species are related
104 PROTEINS | The Protein Chemistry of Cereal Grains

Albumin

Globulin

Prolamin

Glutenin

Ile Leu Lys Met Cys Phe Tyr Thr Trp Val Arg His Ala Asp Glu Gly Pro Ser NH3

Figure 8 Amino acid composition of the Osborne fractions of wheat.

Table 3 The amino acid compositions of whole grains of several the cereal endosperm have been given the specific names
cereals, expressed as grams per 16 g of nitrogen according to each cereal genus, as listed in Figure 9.

Amino acid Wheat Rice Corn Barley Oats

Alanine 3.6 6.0 7.5 4.0 4.5 The Proteins of Wheat, Rye, and Triticale
Arginine 4.6 8.3 4.2 4.7 6.3
Aspartic acid 4.9 10.3 6.3 5.7 7.7 Wheat and rye are closely related; triticale is a man-made
Cysteine 2.5 1.1 1.6 2.3 2.7 hybrid between wheat and rye. The genetic control of the
Glutamic acid 29.9 20.6 18.9 23.6 20.9 storage proteins of these cereals is detailed in the specific
Glycine 3.9 5.0 3.7 3.9 4.7 articles on wheat, rye, and triticale. Their protein compositions
Histidine 2.3 2.5 2.7 2.1 2.1 are thus similar to a limited extent, but the dough-forming
Isoleucine 3.3 3.8 3.7 3.6 3.8
quality of rye and triticale is much poorer than that of wheat.
Leucine 6.7 8.2 12.5 6.7 7.3
The popularity of rye-based foods is not so widespread as that
Lysine 2.9 3.8 2.7 3.5 3.7
Methionine 1.5 2.3 1.9 1.7 1.7 of foods made from wheat. Rye is especially popular in Eastern
Phenylalanine 4.5 5.2 4.9 5.1 5.0 European and Scandinavian countries. Baked products are
Proline 9.9 4.7 8.9 10.9 5.2 commonly crispbread and varieties of bread, including
Serine 4.6 5.4 5.0 4.0 4.7 pumpernickel.
Threonine 2.9 3.9 3.6 3.3 3.3 Whereas true rye bread is made from 100% rye, it is
Tryptophan 1.1 1.2 0.6 1.5 1.3 common for rye bread to be made from a blend of rye and
Tyrosine 3.0 3.5 3.8 3.1 3.0 wheat flours, the contribution of the wheat gluten being
Valine 4.4 5.5 4.8 5.0 5.1 needed to provide the baking quality that is lacking in the rye
flour. The poorer dough-forming characteristics of rye gluten
are largely due to the lower level of the glutelin-type protein,
combined with the more hydrophilic (water-loving) properties
of the rye secalins, compared with the corresponding classes of
genetically, these genetic similarities and distinctions are protein in wheat.
reflected directly in the amino acid sequences of their storage The water-soluble proteins of rye contain the albumin and
proteins. Both the relevant gene sequences and the correspond- globulin classes of protein, similar to those in wheat, but for
ing amino acid sequences also reflect the taxonomic relation- rye, there is generally a higher level of amylase enzymes than in
ships among the cereal species, as is illustrated in Figure 9. To a wheat because of the greater susceptibility of rye to preharvest
large extent, these differences explain how the storage proteins sprouting and the consequent production of starch-degrading
of the respective cereal grains make their distinct contributions enzymes.
with respect to their food uses. The essential amino acid composition of rye is slightly
The most obvious example is wheat, whose gluten-forming better than that of wheat, because of the lower content of
storage protein is unique among the cereals in suiting wheat prolamin-type proteins in rye (Figure 9). The properties of
flour for bread making. Furthermore, the proteins of the rice triticale are intermediate between those of rye and wheat,
grain contribute to its eating quality. The storage proteins of reflecting the genetic origins of triticale.
 PROTEINS | The Protein Chemistry of Cereal Grains 105

Globulins Prolamins

7S 11/12S Other prolamins The prolamin superfamily

Maize, oats, Wheat Oats Rice Maize Sorghum Rice Maize Oats Triticeae:
rice, barley, barley, wheat,
rye, wheat rye

7S globulin T riticin Globulin Glutelin Kaffirin Avenin

16 kDa 10 kDa 13 kDa

-Zein -Zein -Zein -Zein

HMW S-rich S-poor

Embryo,
aleurone Endosperm

Figure 9 Storage protein classes in cereals. Adapted from Shewry, P. R., Tatham, A. S., and Halford, N. G., 1999. The prolamins of the Triticeae. In:
Shewry, P. R. and Casey, R. (Ed.), Seed Proteins. Kluwer Academic, The Netherlands, pp. 3578.

Table 4 Classes of barley proteins (hordeins) and corresponding loci

Hordein class Loci Gene copy number Proteins present in the cv Carlsberg MW (kDa) Corresponding wheat prolamin

B Hor 2 2030 10 3645 LMW glutenins

C Hor 1 2030 9 5060 o-Gliadins
D Hor 3 1 1 105 HMW glutenins
g Hor 5 Low 7 3645 g-Gliadins

The Proteins of the Barley Grain
The protein concentration of cultivated barley ranges from 9%
to 12%. For barley, the protein content is generally calculated
as total nitrogen content multiplied by the factor of 6.25. The
protein content of hull-less (naked) barley genotypes is gener-
ally higher (even up to 18%) because of the absence of the
outer hull layers (the glumes the palea and lemma), which
contain relatively little protein.
The production of the enzymes of germination is an impor-
tant part of a good malting barley, but paradoxically, high
protein content (e.g., over 11%) is not necessarily desirable
for the best malting quality. This is because the ultimate test of
malting quality is the extent of extract produced during the
mashing process. The amount of extract determines how much
beer can be produced from a given amount of malt. Extract
represents the amount of soluble fermentable sugars from the
hydrolysis of starch, resulting from the combination of very
active amylases and high starch content. For this reason, a low
protein content (i.e., less than 8%) is also undesirable, as it
may have the consequence of inadequate enzyme production.
Consumption for human food is a traditional use of barley,
but this now accounts for only 5% of total barley consumption
worldwide. Feed use predominates, amounting to 75% of
barley use. The amino acid composition of barley does not
provide a consistently better balance nutritionally than other
cereal grains, still being deficient in lysine, methionine, and
threonine. Nevertheless, it is a popular source of protein and
energy in feedstuffs, in combination with other sources of
energy. High-lysine barley genotypes have been developed,
analogous to those in maize, the first of these being the
Hyproly line.
The hordein polypeptides show considerable polymor-
phism, like the prolamins of wheat and rye. A large proportion
of the barley prolamins are disulfide-linked (both intra- and
interchain linkages) with naturally occurring polymers of high
molecular weight. Unlike the glutenin polymers of wheat,
these are not desirable for the processing of barley. Because
they generate problems in the malting process and down-
stream, they are likely to produce haze in the final product
(i.e., beer) if they have not been adequately hydrolyzed during
processing.
The hordein polypeptides are classified into groups (A, B, C,
and D hordeins) according to their decreasing mobilities dur-
ing SDS gel electrophoresis. The C hordeins have analogy to
the omega-gliadins of wheat, having a general absence of the
sulfur-containing amino acids. They are coded by genes (locus
Hor 1) on the short arm of barley chromosome 1H, as indi-
cated in Table 4. The genes (Hor 2 and Hor 5) for the sulfur-rich
B and gamma-hordeins are also coded on chromosome 1H.
The D hordeins, which occur as disulfide-linked polymers, are
analogous to the HMV subunits of wheat glutenin; like them,
106 PROTEINS | The Protein Chemistry of Cereal Grains
their genes (Hor 3) are located on the long arm of the corre-
sponding barley chromosome (1H).
The Proteins of the Rice Grain
The proteins of rice are a contrast to those of wheat and barley,
being predominantly of the globulin class. As a protein source,
milled rice contains the lowest amount of protein (about 5%)
among the major cereals. Moreover, this protein is not easily
digestible by humans and monogastric animals. However,
compared with that of other cereal proteins, the overall
amino acid composition of rice protein is significantly more
balanced due to the relatively higher level of lysine content.
This unusual amino acid composition results because rice is
one of the few cultivated plants in which there are significant
levels of globulins and prolamins. These are the two major
classes of storage proteins in the seeds of higher plants. Unlike
other cereals that accumulate prolamins as their primary nitro-
gen reserve, the major storage proteins in rice are the glutelins,
which are homologous at the primary sequence level to the 11S
globulin proteins (Figure 9), a class that is the dominant form
of nitrogen deposition in legumes. Rice prolamin proteins
have a number of characteristics that are different from the
prolamins present in most other cereals.
The most important characteristic of the deposition of
nitrogen in the rice kernel is that there are three kinds of
protein bodies in the rice endosperm, namely, large spherical,
small spherical, and crystalline protein bodies. Each of these is
surrounded by a single continuous membrane. Although the
spherical protein bodies form within vacuoles, the proteins are
synthesized in the endoplasmic reticulum and in the Golgi
apparatus and are then transported to the vacuoles via vesicles.
Removal of the husk (hull) from rough (paddy) rice yields the
kernel. This is composed of the pericarp, seed coat, aleurone,
endosperm, and the germ and is known as brown rice. It has a
protein content of 910%, with a significantly higher nutri-
tional value than the most commonly utilized rice product
white polished (milled) rice, whose protein content is 8%.
The milling process for paddy rice results in 4055% white
milled rice, together with three major by-products, namely,
husks (20%), bran (10%), and brokens (1022%). These by-
products have protein contents of 3%, 17%, and 9%, respec-
tively. The aleurone layer, the tissue with the highest level of
proteins and nutritionally important minor components, is
removed during the rice milling process.
The subaleurone region of the rice grain plays an important
part because of its nutritional value. It is a globulin-rich layer,
being several cell layers thick. Its lysine content is much higher
than that of the proteins located in the rice endosperm. Thus,
rice should be milled as lightly as possible to retain as much as
possible the subaleurone layer.
The albumin fraction of rice is highly heterogeneous, con-
taining biologically important components. It can be separated
into four subfractions, based on the molecular size of its pro-
teins. More than 50 individual polypeptides were observed in
the albumin fraction, using isoelectric focussing. As for cereal
grains in general, the albumins are mainly enzymes and
enzyme inhibitors.
Like the prolamins from other cereals, rice prolamins are
readily soluble only in alcohol/water mixtures. However, the
classical extraction procedure had not been useful in early
studies. More recently, these estimates of prolamin content
have been shown to be low. Using 55% propan-L-ol and reduc-
ing agents, higher yields of prolamins (2025% of the total
protein) have been obtained. Rice prolamins are 1016 kDa in
molecular weight. Thus, their molecular size range is signifi-
cantly smaller than the prolamins of other cereals. They are
highly variable between rice cultivars, based on their electro-
phoretic and isoelectric focussing fingerprints.
According to the Osborne classification system, the least
soluble proteins of the rice grain are oligomeric globulins
representing up to 80% of the total protein fraction
(Figure 9). They have been difficult to study because of their
general insolubility in all solvents except dilute alkali, due to
their high molecular weight and their heterogeneity. When
analyzed by gel filtration chromatography, the purified globu-
lin subunits were resolved into three subunits, linked by disul-
fide bonds, which varied in stoichiometry depending on the
report and on the rice variety used.
Despite their general insolubility, rice glutelins are homol-
ogous in structure to 11S globulins (Figure 9). The structural
relationship is evident when comparing the N-terminal amino
acid sequences and using immunologic methods. The extent of
homology between the rice glutelin and legume 11S globulins
is 3035%. Structural similarity is evident in the primary
sequences of homology to a motif common in wheat, rye,
barley, and maize prolamins.
The Proteins of Maize
Maize (or corn), often classed in world trade as one of the
coarse grains, is an important source of protein in the diets of
many people in countries such as Africa and South America. In
addition, maize is used extensively as a feed grain and in a
range of industrial processes. The process of corn starch man-
ufacture produces the grain protein as a by-product. It is some-
times referred to as corn gluten in the trade, but this name is
inappropriate, because maize protein has no relationship to
the gluten of wheat and is safe for consumption by those
susceptible to celiac disease.
Like the other cereal grains, maize protein is slightly defi-
cient in lysine. This limits its utilization as an animal feed. This
nutritional imbalance is largely due to the low lysine content of
zein, the major group of maize proteins (Figure 9). The
development of the opaque-2 and floury-2 mutations has
led to the development of maize genotypes with higher levels
of lysine, due to a lower content of zein and more of the
nutritionally superior albumin type of protein. Although
these new maize types offer nutritional advantages, they carry
a grain yield penalty.
The zein protein of maize can be isolated by extraction of the
crushed endosperm (the starchy part of the grain) with 70%
aqueous ethanol, after breaking disulfide bonds. Maize zein has
been further fractionated into subclasses of proteins, designated
alpha-, beta-, gamma-, and delta-zeins. The genes for their syn-
thesis have been mapped to maize chromosomes 6 and 7.

The Proteins of Sorghum
The major storage proteins of sorghum, termed kafirins
(Figure 9), are fractionated by a procedure similar to that for
the zeins of maize. Alpha-kafirin, composing about 80% of the
overall kafirin content, can be further subdivided into two
proteins with molecular weights of about 23 000 and
25 000 Da. Three beta-kafirins and one gamma-kafirin have
been identified, all of them having molecular weights in the
range of 1628 kDa. Like maize, high-lysine mutants of sor-
ghum have been developed.
Future Prospects
Our current understanding of grain protein chemistry contrasts
strongly with the rudimentary knowledge of the early twentieth
century. That knowledge has progressed from the recognition
of only a few distinct grain proteins to fractionating and char-
acterizing thousands of proteins from all the tissues of the
cereal grain the endosperm, germ, scutellum, and bran.
This knowledge permits more efficient selection for improved
processing quality in the breeding of new varieties. The knowl-
edge is also being applied to testing for protein quality after the
grain is harvested, so that grains of specific quality can be
segregated for appropriate marketing and processing.
Knowledge at the chemical level is now complementing
new genetic insights, so that breeders and molecular geneticists
can employ conventional and novel methods to create quality
types that have previously not been possible varieties with
new protein functionality. Thus, the future holds great promise
for grains that will be processed more efficiently with nutri-
tional advantages. For example, it has been possible to incor-
porate purified gluten proteins into rice dough, thus to
monitor their effects on the mixing properties.
The protein content of the cereal grains has always been
important for marketing and processing. The generally desir-
able trait of high-protein concentration has often been
inversely proportional to grain yield. Improvements in genet-
ics and agronomic practice have tended to reduce grain pro-
tein levels, because high yield has been a common goal for
farmers.
The coming decades will bring a new factor that will further
erode protein levels the rising concentrations of carbon
dioxide in the atmosphere. For whatever reason, it is inevitable
that CO2 levels will continue to rise, although the rate of
increase is difficult to predict. Many experiments have been
conducted to simulate effects of future levels of CO2. They have
consistently indicated increased yields of grain and biomass,
together with decreased levels of grain protein content. The
higher atmospheric concentration of CO2 acts as a carbon
fertilizer, with consequent increases in starch synthesis but
little change in protein synthesis. There will thus be a need to
rethink the role of protein in cereal grains if it will gradually
become lower in concentration.
At the manufacturing level, there is increasing interest in pro-
viding greater protein content for cereal-based snacks and break-
fast foods. Cereal proteins are also undergoing texturization and
extrusion with the aim of creating synthetic meat products.
PROTEINS | The Protein Chemistry of Cereal Grains 107
Exercises for Revision
How did the early chemists views on protein chemistry
differ from our present knowledge? Why were they unaware
of the great heterogeneity of grain proteins?
What are the aspects of protein composition that explain
the great differences between the various proteins in their
properties?
Prepare a table of differences and similarities between the
major cereal species with respect to their protein
compositions.
What are the nutritional deficiencies of cereal grain proteins
with respect to the essential amino acids? Describe the genetic
achievements that have partially overcome these deficiencies.
Exercises for Readers to Explore the Topic Further
Examine the taxonomic relationships between the major
cereal species, as provided in this encyclopedia. Compare
the associations between the proteins of these grains in
Figure 8. Look at the genus names to see the origins of
the grain-specific protein names.
Obtain the amino acid compositions of the grain proteins
for the major cereals. What are the main differences? How
would these differences account for functional and nutri-
tional differences between these grain species?
Contrast the differences in function for an enzyme (to
catalyze a specific reaction) versus a storage protein
(a source of amino acids for the new plant). Why would
both types of protein be produced under the same strict
genetic control?
Compare the various methods of determining the amount
of protein in a grain or flour sample (Table 2 and else-
where). Which would you expect to be the most accurate?
See also: Genetics of Grains: Maize: Other Maize Mutants; Maize:
Quality Protein Maize; Grain Marketing and Grading: Identification
of Varieties of Food Grains; Proteins: Protein Synthesis and
Deposition; Proteomic Analysis Relevant to Cereal Grains; The
Enzymes Associated with Food Grains and Their Functions in
Processing; The Basics: Taxonomic Classification of Grain Species;
The Grain Crops: An Overview; The Cereal Grains: An Overview of
the Family of Cereal Grains Prominent in World Agriculture; Wheat
Processing: The Gluten Proteins of the Wheat Grain in Relation to
Flour Quality; Wheat-Based Foods: Cereals: Overview of Uses:
Accent on Wheat Grain.
Further Reading
Bailey HC (1941) A translation of Beccaris lecture Concerning Grain (1729). Cereal
Chem. 18: 555.
Bushuk W (ed.) (2001) Rye: Production, Chemistry and Technology, second ed. St.
Paul, MN: American Association of Cereal Chemists.
Dendy DAV (ed.) (1995) Sorghum and Millets: Chemistry and Technology St. Paul, MN:
American Association of Cereal Chemists.
Hartley H (1951) Origin of the word protein. Nature 168: 244.
Henry RJ and Kettlewell PS (1996) Cereal Grain Quality. London: Chapman and Hall.
Islam S, Ma W, Yan G, Bekes F, and Appels R (2011) Modifying processing and health
attributes of wheat bread through changes in composition, genetics and breeding.
108 PROTEINS | The Protein Chemistry of Cereal Grains
In: Cauvain SP (ed.) Bread making. Improving quality, second ed. Boston,
New York: CRC Press.
Juhasz A, Bekes F, and Wrigley CW (2014) Wheat proteins. In: Ustunol Z (ed.) Applied
Food Protein Chemistry, Chapter 11, pp. 219304. West Sussex, UK:
Wiley-Blackwell, John Wiley & Sons.
Juliano BO (ed.) (1985) Rice: Chemistry and Technology, second ed. St. Paul, MN:
American Association of Cereal Chemists.
Kent NL (1975) Technology of Cereals, with Special Reference to Wheat, second ed.
Oxford, UK: Pergamon.
Lasztity R (1999) Cereal Chemistry. Budapest, Hungary: Akademiai Konyvkiado. Wrigley CW (2012) Proteins the basis of life. Teach. Sci. 58: 5659.
McGregor AW and Bhatty RS (eds.) (1993) Barley: Chemistry and Technology. St. Paul,
MN: American Association of Cereal Chemists.
Mertz ET (ed.) (1992) Quality Protein Maize. St. Paul, MN: AACC International.
Pomeranz Y (ed.) (1988) Wheat: Chemistry and Technology, third ed. St. Paul, MN:
American Association of Cereal Chemists.
Seneweera S, Fernando N, Panozzo J, Tausz M, Norton RM, Fitzgerald GJ, Myers S, and
Nicolas ME (2014) Intra-specific variation of wheat grain quality in response to
elevated [CO2] at two sowing times under rain-fed and irrigation treatments.
J. Cereal Sci. 59(2): 137144.
Shewry PR and Casey R (eds.). (1999) Seed Proteins. The Netherlands: Kluwer
Academic.
Shewry PR and Miflin BO (1985) Seed storage proteins. In: Pomeranz Y (ed.) Advances
in Cereal Science and Technology vol. 7, pp. 183. St. Paul, MN: American
Association of Cereal Chemists.
Sikorski ZE (2001) The Chemical and Functional Properties of Food Proteins.
Lancaster, PA: Technomic Publishing.
Watson SA and Ramsted PE (eds.) (1987) Corn: Chemistry and Technology St. Paul,
MN: American Association of Cereal Chemists.
Webster FH (ed.) (1985) Oats: Chemistry and Technology. St. Paul, MN: American
Association of Cereal Chemists.
Wrigley CW and Batey IL (eds.) (2010) Cereal Grains: Assessing and Managing Quality
Oxford, UK: Woodhead Publishing Ltd.
Relevant Websites
http://www.aaccnet.org American Association of Cereal Chemists.
http://www.campden.co.uk Campden & Chorleywood Food Research Association.
http://www.icc.or.at International Association for Cereal Science and Technology.
http://www.usda.gov United States Department of Agriculture.
http://www.wheat.pw.usda.gov Grain genes.
 The Protein Chemistry of Dicotyledonous Grains
PR Shewry, Rothamsted Research, Harpenden, UK; University of Reading, UK
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The major types of storage proteins present in seeds of
dicotyledonous (noncereal) crops are globulins and
albumins.
Two types of storage globulin are recognized with sedimen-
tation coefficient S20,w values of
7 and 11.
7S and 11S globulins have related structures and belong to
the cupin superfamily of plant proteins.
2S albumin storage proteins are related to other types of
small sulfur-rich proteins and belong to the prolamin
superfamily, which includes cereal seed storage proteins.
Learning Objectives
To understand storage protein composition and properties
in dicotyledonous crops in relation to evolutionary rela-
tionships and biological roles.
Introduction
The protein contents of dicotyledonous grain crops vary
widely, from less than 20% of the dry weight in the pseudocer-
eals to
50% in some lines of soybean (Table 1). Extraction
and separation of total protein fractions show that all com-
prise highly complex mixtures, with many components being
present in small amounts. However, in all cases, a small num-
ber of components are present in substantial amounts: these
correspond to the seed storage proteins and usually account for
half or more of the total seed proteins.
Although specialized storage protein types occur in some
species, for example, lectins in some legume seeds, most of the
storage proteins present in dicotyledonous seeds fall into three
groups, which are defined on their solubility and S20,w values
(sedimentation coefficients determined by sedimentation
equilibrium ultracentrifugation), which are a measure of their
molecular masses. The characteristics of these three types of
proteins are discussed, followed by brief descriptions of the
storage proteins present in the major dicotyledonous species
covered in this encyclopedia.
Types of Storage Protein
Globulins
Globulins are defined as proteins that are soluble in dilute salt
solutions, usually taken as 0.51.0 M NaCl. Storage globulins
with S20,w values of
7 and 11 are widely distributed in dicot-
yledonous plants. However, they have been most widely stud-
ied from legume seeds. Hence, 7S globulins are sometimes
called vicilins after the Viciae tribe of legumes, and 11S
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00105-4
globulins are called legumins after the family Leguminosae.
The individual types of 7S and 11S globulins in different crops
are also often given specific trivial names, as listed in Table 1.
It has long been recognized that members of these two
protein groups also vary in their precise S20,w values, with
legumin-like globulins often having values of 12 or more
(e.g., the 13S lupin globulin) and vicilin-like globulins

8 (e.g., in peanut). Furthermore, not all 11S globulins are
readily soluble in dilute saline, with notable exceptions being
11S storage globulins of cereals, which are either insoluble in
saline (rice) or only soluble at higher concentrations
(0.81.0 M) of NaCl (oats). Consequently, the term 11S is
used in this article to describe storage proteins with a
legumin-like structure and the term 7S to describe vicilin-like
proteins.
7S globulins
7S globulins typically have molecular masses of

150200 kDa and have a trimeric structure comprising
three nonidentical subunits of mass
5070 kDa. The sub-
units contain no cysteine residues and hence do not form
disulfide bonds, and noncovalent forces stabilize the trimeric
proteins. However, this simple structure may be modified after
protein synthesis in two ways. First, glycosylation (addition of
oligosaccharides) may occur on one or two asparagine residues
in one or more subunits of the trimer. This results in an
increase in mass of the subunits and lower mobility when
separated by sodium dodecylsulfate polyacrylamide gel elec-
trophoresis (SDS-PAGE) with partial glycosylation resulting in
additional heterogeneity.
Second, in some species (e.g., pea and field bean), some,
but not all, 7S globulin subunits are proteolytically processed
at one or two sites, giving rise to polypeptide chains ranging in
mass from processed products of
12 kDa in pea and 18 kDa
in bean to unprocessed subunits of
50 kDa. The processed
products remain associated in the trimeric complexes until
released by denaturation. The role of this proteolytic proces-
sing is not known and it does not occur in some species such as
soybean and common or navy bean (Phaseolus vulgaris).
Although typically trimeric in structure, the 7S globulins may
undergo reversible aggregation into hexamers depending on
ionic strength.
11S globulins
11S globulins typically have masses of
300400 kDa and are
hexamers of six nonidentical subunits. Each subunit is synthe-
sized as a precursor protein that is proteolytically processed to
release acidic (also called A or a) and basic (B or b) polypeptide
chains, which correspond to the N-terminal and C-terminal
parts of the precursor protein, respectively. The masses of the
acidic chains vary in the range of 2060 kDa but are typically
109
110 PROTEINS | The Protein Chemistry of Dicotyledonous Grains

Table 1 Protein contents and types of storage proteins in major Table 2 Polypeptide molecular masses (Mr  103) of 7S and 11S
legume, oilseed, and pseudocereal crops globulins present in species

Protein content 7S 11S acidic chain 11S basic chain

Species (% DW)a Storage proteins
Legumes
Legumes Soybean 50, 80 1040 20
Soybean 3040 12S glycinin, 7S b-conglycinin, 2S Pea 1250, 2540 2122
albumin 70
Pea 25 12S legumin, 7S vicilin/convicilin Peanut 2375 1448 21
2S PA1 Lupin 1560 2563 21
Peanut 1636 13S arachin, 8S conarachin, 2S Oilseeds
albumin Oilseed rape/ 3040 20
Lupin 2642 1112S conglutin a, 7S conglutin canola
b, 2S conglutin d Sunflower 3234 2227
Oilseeds Pseudocereals
Oilseed rape/ 3644 12S cruciferin, 2S napin Quinoa 3239 2223
canola Amaranth 1590 3037 1827
Linseed/ 2024 12S globulin Buckwheat 1666 3243 2023
linola
Sunflower 1520 12S helianthinin, 2S albumin The terms 7S and 11S refer to vicilin-like and legumin-like proteins, respectively, and
Safflower 15 12S carmin, 2S albumin do not imply that all of the proteins listed have these precise S20,w values. In fact, the
Cottonseed 1622 11S globulin, 7S globulin, 2S true values vary (
78S and 1113S, respectively). It should also be noted that widely
albumin different values for polypeptide masses have been reported and those presented here
Pseudocereals have been selected by the author. The 7S polypeptides include whole subunits and
Quinoa 15 11S chenopodin, 2S albumin chains released by proteolysis. The acidic and basic 11S chains are generated by
Amaranth 1318 13S amaranthin, 8S proteolysis of a single precursor subunit.
conamaranthin, 2S albumin
Buckwheat 13.5 11S globulin, 8S globulin, 2S
albumin Prokaryotic
protogermin
The S20,w values listed are only approximate with a range of values having been reported Slime Plants protein
for some components. molds
a
% Dry weight.
Spherulins

40 kDa, while the basic chains usually have masses of

20 kDa. Thus, a typical 11S globulin comprises six subunits Duplicated ancestor
of mass
60 kDa, each consisting of 40 and 20 kDa polypep- Germins- and
germin-like
tide chains. The basic and acidic chains are covalently linked by
proteins
a single disulfide bond formed in the precursor protein.
Whereas the 7S globulins are rapidly assembled into their
Fern vicilin-like
mature trimeric structure, the 11S globulin precursors are not protein
immediately assembled into their mature hexameric structure
but into an intermediate trimer with the hexamer forming only
after proteolytic processing occurs to release the basic and acidic basic N C
acidic chains. 11S globulins 7S globulins
The polypeptide (i.e., subunit and chain) masses of the 7S (legumins) (vicilins)
and 11S globulins discussed in the succeeding text are summa-
Figure 1 Hypothetical pathway for the evolution of germins
rized in Table 2. and globulin storage proteins from an ancestral protogermin protein.
Reproduced with permission from Shewry, P.R., Jenkins, J.,
Beaudoin, F., Mills, E.N.C., 2003. The classification, functions and
Structures of 7S and 11S Globulins
evolutionary relationships of plant proteins in relation to food allergens.
Simple comparisons of the amino acid sequences of 7S and In: Mills, E.N.C., Shewry, P.R. (Eds.), Plant Food Allergens. Blackwell
11S globulins show little or no evidence of homology, but Science, Oxford, UK.
more sophisticated comparisons (in which the properties of
the amino acids and their propensity to form similar structures ancestral protein that initially underwent a duplication to
are also considered) do indicate some relatedness, particularly give two homologous domains, which diverged to form the
between the C-terminal parts of the 7S subunit and the basic 7S N-terminus/11S basic chain and 7S C-terminus/11S acidic
chains of 11S subunits. Similar comparisons also show inter- chain, as shown in Figure 1.
nal homology between the N- and C-terminal parts of the 7S Comparisons of the three-dimensional structures of 7S
subunits. These observations, and the more detailed structural globulin trimers (phaseolin from common bean, canavalin
comparisons discussed in the succeeding text, indicate that the from jack bean, and b-conglycinin from soybean) and the
7S and 11S globulin subunits have evolved from a single intermediate trimer of proglycinin (the 11S globulin of
 PROTEINS | The Protein Chemistry of Dicotyledonous Grains 111

-Helical -Helical
domain 4 domain 2

Domain 3
Domain 1
Figure 2 Structure of the 7S globulin canavalin from jack bean (Canavalia ensiformis) seeds. The protein is trimeric, with a single subunit
shown in lighter gray. Domains 1 and 3 are b-barrels and domains 2 and 4 a-helical. Reproduced with permission from Shewry, P.R., Jenkins, J.,
Beaudoin, F., Mills, E.N.C., 2003. The classification, functions and evolutionary relationships of plant proteins in relation to food allergens.
In: Mills, E.N.C., Shewry, P.R. (Eds.), Plant Food Allergens. Blackwell Science, Oxford, UK.

soybean) confirm this scheme and show remarkable conserva-
tion of structure. Each protein consists of three subunits
arranged around a threefold axis (Figure 2), with each subunit
consisting of two closely related structural modules and each
comprising an antiparallel b-barrel domain and an a-helical
domain. The subunits of 7S and 11S globulins have similar
structures, as shown by the overlay in Figure 3, and the mature
hexameric 11S globulin protein is considered to be a dimer of
trimers.
The similarity in the structures of the 7S and 11S globulins
is consistent with the observation that both proteins can exist
in trimeric forms (the mature 7S and intermediate 11S globu-
lins) and in hexameric forms (the mature 11S globulins and
the mature 7S globulins under specific conditions). It is also
consistent with the fact that the 7S and 11S proteins appear to
be stored as a homogeneous mixture in protein bodies in seeds
of legumes and other dicotyledonous species.
A final point of interest is that both 11S and 7S globulins
may be allergenic, notably in soybean and peanut.
Albumins
Albumins are defined as proteins that are soluble in water.
Consequently, albumin fractions extracted from most plant
tissues comprise complex mixtures of largely unrelated com-
ponents. However, the seeds of many dicotyledonous plants
contain a well-defined family of albumin storage proteins with
S20,w values of
2. These 2S albumins are major storage pro-
teins in a number of dicotyledonous crops (e.g., oilseed rape/
canola and sunflower) and minor components in others (e.g.,
soybean). Although they have not yet been identified in some
dicotyledonous crops, this may reflect lack of study rather than
true absence, and they may well prove to be ubiquitous in
dicotyledonous seeds. A preliminary report of the presence
of 2S albumin storage protein in seeds of one monocotyledon-
ous species (yucca) has not been confirmed by more detailed
studies, but they are known to be structurally related to several
other families of low-molecular-mass sulfur-rich proteins
including nonspecific lipid transfer proteins and trypsin/a-
amylase inhibitors of cereal grain.
A typical 2S albumin consists of two subunits of
Mr
810 kDa and 45 kDa, which are associated by two
disulfide bonds. However, these two subunits are initially syn-
thesized as a single precursor polypeptide, and hence, by anal-
ogy with 7S and 11S storage globulins, they should perhaps be
called polypeptide chains. Nevertheless, the term subunit is
widely used and will be retained here.
The proteolytic processing to release the 2S albumin sub-
units may result in the loss of short peptide sequences from the
N- and C-termini of the proteins and from between the two
subunits (linker peptides). Furthermore, this proteolysis only
occurs after the proproteins are folded and four disulfide
112 PROTEINS | The Protein Chemistry of Dicotyledonous Grains

Figure 3 Overlay of the backbone structures of single subunits of proglycinin (11S globulin precursor) from soybean (black) and phaseolin
(7S globulin) of bean (gray). Reproduced with permission from Shewry, P.R., Jenkins, J., Beaudoin, F., Mills, E.N.C., 2003. The classification, functions
and evolutionary relationships of plant proteins in relation to food allergens. In: Mills, E.N.C., Shewry, P.R. (Eds.), Plant Food Allergens.
Blackwell Science, Oxford, UK.

Signal peptide
removed
cotranslationally

21 157
NH2 S A B C COOH
Small Large subunit
subunit
N-terminal C-terminal
prosequence Linker prosequence
peptide
Peptides removed by
proteolysis in vacuole
Figure 4 Schematic sequence of the precursor protein of napin (the 2S albumin of oilseed rape) indicating peptides removed by proteolysis and
the large and small subunits of the mature protein.

bonds formed between eight highly conserved cysteine resi-

dues. Two of these become interchain bonds linking the two
subunits and two intrachain bonds within the large subunits.
This processing is shown schematically in Figure 4. The mature
proteins have a compact tightly folded structure comprising
five a-helices in a right-handed fold (Figure 5).
The vast majority of 2S albumins have two-chain structures
similar to that described earlier. However, single-chain forms
(i.e., without proteolytic processing into two chains) also
occur, notably in sunflower.
Of particular interest in relation to grain nutritional quality
is the presence in some species of 2S albumins that are rich in
methionine or, less commonly, cysteine, the two sulfur-
containing amino acids that are frequently limiting in legume
and other dicotyledonous seeds. These albumins have been
studied in most detail in Brazil nut and sunflower. Although
Figure 5 Schematic structure of the 2S albumin from oilseed rape.
they were initially considered to provide an attractive target for Alpha helices are shown as cylinders and numbered 15. Reproduced
use in genetic engineering to increase seed sulfur content, it is with permission from Shewry, P.R., Jenkins, J., Beaudoin, F., Mills,
now known that both are allergenic, as are 2S albumins from a E.N.C., 2003. The classification, functions and evolutionary relationships
number of other species (e.g., mustards, castor bean, walnut, of plant proteins in relation to food allergens. In: Mills, E.N.C., Shewry,
sesame seeds, and peanut). P.R. (Eds.), Plant Food Allergens. Blackwell Science, Oxford, UK.
 PROTEINS | The Protein Chemistry of Dicotyledonous Grains 113

Storage Proteins of Legumes the 7S conglutin b, with slightly less conglutin a (11S globulin)
and
25% of the 2S albumin conglutin d. Conglutin a has a
Many leguminous species store 7S globulin, 11S globulin, and
mass of
336 kDa and readily dissociates to a trimeric form
2S albumin proteins, but the proportions of these vary and
(7.1S, Mr 170 kDa) at low ionic strength or high pH (8.8). It
exceptions do occur. For example, Phaseolus contains 7S but
comprises acidic chain varying in Mr (within and between
not 11S globulins. Additional types of storage protein also
species) from 25 to 63 kDa linked to basic chains of
occur in some species, including various lectins and protease
Mr
21 kDa. Conglutin a is unusual among 11S legumin-
inhibitors, which can have negative impacts on nutritional
type globulins in that it is glycosylated on a single site. Con-
quality.
glutin b comprises polypeptides that range in mass between
Soybean is the most widely studied legume species. It con-
and within species from
15 to 60 kDa.
tains major groups of 11S (glycinin) and 7S (b-conglycinin)
Little detailed work has been carried out on the other
globulins, in ratios ranging from
1:1 to 6:1 depending on the
legumes/pulses discussed in this article. Both chickpea and
genotype, with only small amounts of 2S albumins. In addi-
lentil contain 11S and 7S storage globulins (in a ratio
tion, a number of other proteins can each account for up
of
6:1 in chickpea). In addition, chickpea contains a major
to
5% of the total protein in some lines, notably Kunitz-
albumin of Mr
20 kDa, which comprises subunits with Mr 10
and BowmanBirk-type protease inhibitors, lectins, urease,
and 12 kDa.
and lipoxygenase. Sedimentation coefficients and masses for
Adzuki bean is rich in 7S globulins with major components
glycinin have been reported to range from 11.8S to 13.1S and
of Mr
55 kDa, but small amounts of 11S globulins and 2S
from 30 to 380 kDa and for b-conglycinin from 6.7S to 8.0S
albumins are also present.
and from 105 to 330 kDa.
Glycinin has a typical hexameric 11S globulin structure,
with subunits comprising basic chains of Mr
20 kDa linked Storage Proteins of Oilseeds
to acidic chains varying in mass from
10 to 40 kDa. Glycinin
The two most intensively studied oilseeds are sunflower and
is also unusual in that the hexamer can undergo reversible
oilseed rape (canola). Both contain predominantly 11S-type
dissociation into trimers, depending on the ionic strength.
globulins with some 2S albumins.
b-Conglycinin comprises various combinations of three
The 12S globulins (cruciferin) of oilseed rape account
major subunits a (Mr
80 kDa), a0 (Mr
80 kDa), and b
for
40% of the total seed proteins and have Mr of
(Mr
50 kDa) resulting in seven major forms. Both 7S and
30360 kDa. They have a typical hexameric structure with
12S globulins of soybean are also thought to be allergenic.
acidic chains of Mr 3040 kDa linked to basic chains of Mr
Pea seeds contain legumins (12S to 13S, Mr 330420 kDa)
20 kDa. The 2S albumins of oilseed rape (napins) also have a
and vicilins (7S to 8S, Mr 150190 kDa). The legumin subunits
typical structure. The major napins have masses of
14.5 kDa
consist of acidic chains of Mr
25 kDa and 40 kDa linked to
(subunit Mr of
9 and 4 kDa) but minor forms of
basic chains of Mr
20 kDa. The 7S vicilins comprise subunits
Mr
12.5 kDa also occur. Napin processing includes the loss
of Mr
50 kDa, but proteolysis at one or two sites and glyco-
of N-terminal, C-terminal, and linker peptides, as discussed
sylation at a single site may occur, resulting in polypeptide
earlier.
chains varying in Mr (1250 kDa). Larger subunits of
Sunflower, a member of the family Compositae, contains
Mr
70 kDa also occur, which form a second trimeric 7S pro-
12S globulins (helianthinin) and 2S albumins in a ratio
tein called convicilin. Pea seeds also contain two major albu-
of
2:1. The Mr 300 kDa helianthinin comprises acidic chains
mins, PA1 and PA2, of which the former may be a 2S storage
of Mr 3234 kDa (a and a0 chains) linked to basic chains of
albumin. PA1 accounts for
10% of the total seed protein but
Mr 2227 kDa (b chains). Sunflower 2S albumins comprise
half of the total sulfur, in the form of cysteine (
11 mol.%).

1113 components with masses of 1018 kDa. All are single-
The dominant peanut proteins are arachin (13S14S, Mr
chain proteins (i.e., they lack a proteolytic cleavage site
330400 kDa) and conarachin (8S to 9S, Mr 140190 kDa).
between the putative large and small subunits), but most are
The subunits of arachin comprise acidic chains of
synthesized as pairs, two albumins being released from a single
Mr
1448 kDa linked to basic chains of Mr
21.4 kDa,
precursor protein by proteolysis. One sunflower albumin that
whereas conarachin comprises polypeptide chains ranging in
is synthesized from a separate precursor protein is called SFA8.
Mr from
23 to 75 kDa. It is probable that both large and
This contains 16 methionine and 8 cysteine residues in a
typical conarachin subunits are present, as in soybean and
mature protein of 103 residues (i.e., a total of 23 mol.% of
pea, with proteolytic processing contributing to the polymor-
sulfur-containing amino acids). Safflower, a related member of
phism. Both proteins also exhibit complex association/disso-
the family Compositae, also appears to contain major groups
ciation behavior, depending on the ionic strength. The major
of 12S globulin (called carmin) and 2S albumin storage
peanut allergen Ara h 1 is a glycosylated 7S globulin with
proteins, but little detailed work has been carried out.
Mr
63.5 kDa, whereas Ara h 3 is an 11S globulin.
Cottonseed is unusual among nonleguminous dicotyle-
Most of the knowledge of peanut albumins comes from the
donous species in containing 7S globulin storage proteins as
analysis of genomic clones and is related to their role as aller-
well as 11S globulins and 2S albumins, with most of the
gens (Ara h 2, 4, 6, and 7). They show high homology to 2S
knowledge being based on DNA-derived sequences with little
albumins from other legumes (soybean and lupin) and are
direct analysis of the seed proteins. Linseed (and linola) has
presumed to have a typical heterodimeric structure.
not been widely studied but the major seed protein is reported
The salt-soluble (i.e., albumin and globulin) proteins of
to be a 12.7S globulin of Mr
250300 kDa. SDS-PAGE
lupins are called conglutins. The major component is usually
showed six nonidentical subunits.
114 PROTEINS | The Protein Chemistry of Dicotyledonous Grains
Storage Proteins of Pseudocereals
Pseudocereals are not true cereals but three dicotyledonous
crops that produce small grain-like seeds. Consequently, their
seed proteins are similar to those of other dicotyledonous
plants rather than related to cereal seed storage proteins.
Seeds of amaranth (Amaranthus spp., Amaranthaceae)
contain 13S legumin-type globulins (amaranthin) and 8S
vicilin-type globulins (S20,w value of 8.6) (conamaranthin).
In addition, albumin proteins with S20,w values of
2 are
present, with SDS-PAGE showing major bands of Mr 9 kDa
and 4 kDa. This is consistent with the presence of typical 2S
albumins. Two methionine-rich proteins with Mr
18 kDa
have also been reported. Quinoa (Chenopodium quinoa,
Chenopodiaceae) seeds also contain 11S globulins (chenopo-
din), of Mr
320 kDa, and 2S albumins, the latter being rich in
cysteine (15.6 mol.%) but low in methionine.
In buckwheat (Fagopyrum esculentum, Polygonaceae), the
major 13S legumin-like globulin is located in the cotyledons
with smaller amounts of putative 8S globulins in the peri-
sperm. Methionine-rich (
9 mol.%) 2S storage albumins are
also present.
Conclusions
The 2S albumins, 7S globulins, and 11S globulins are major
groups of storage proteins that are widely distributed in seeds
of dicotyledonous plants, although their relative amounts vary
and other types of storage proteins also occur in some species.
Consequently, they largely determine the nutritional quality
and processing properties of the seeds. A detailed knowledge of
their structures and properties is therefore essential in order to
elucidate structurefunction relationships. This will facilitate
the optimization of seed quality for traditional and novel
products with improvement using classical genetic or genetic
engineering approaches.
Exercises for Revision
Compare the structures and properties of albumin and
globulin storage proteins.
Compare the contents of albumin and globulin storage
proteins in the major types of dicotyledonous crops:
legumes and pulses, oilseeds, and pseudocereals.
What are the relationships of albumin and globulin storage
proteins to other grain proteins?
Exercises for Readers to Explore the Topic Further
How do the structures and compositions of storage proteins
determine the functional properties of the seeds for food
processing and their role in the human diet?
How can these structures and properties be manipulated
(by breeding or GM technology) to improve their quality
for nutrition and processing and to reduce adverse reactions
in humans (allergies and intolerances)?
See also: Oilseed and Legume Processing: Canola: Processing;
Soybean: Soy Concentrates and Isolates; Proteins: Protein Synthesis
and Deposition; The Protein Chemistry of Cereal Grains; The Basics:
Taxonomic Classification of Grain Species; The Legumes and
Pseudocereals: Amaranth: Overview; Beans: Overview; Buckwheat:
Overview; Chickpea: Overview; Lupin: Overview; Pea: Overview;
Pseudocereals: Overview; Quinoa: Overview; The Oilseeds: Oilseeds:
Overview; Sunflower: Overview; Wheat Processing: The Gluten
Proteins of the Wheat Grain in Relation to Flour Quality; Pulses,
Overview.
Further Reading
Bewley JD and Black M (1999) Seeds Physiology of Development and Germination.
New York: Plenum.
Black M and Bewley JD (2000) Seed Technology and its Biological Basis. Sheffield:
Academic Press.
Boulter D and Croy RRD (1997) The structure and biosynthesis of legume seed storage
proteins: a biological solution to the storage of nitrogen in seeds. Adv. Bot. Res.
27: 184.
Salunkhe DK, Chavan JK, Adsule RN, and Kadam SS (1992) World Oilseeds Chemistry,
Technology, and Utilisation. New York: Van Nostrand Reinhold.
Shewry PR (1995) Plant storage proteins. Biol. Rev. 70: 375426.
Shewry PR and Casey R (eds.) (1999) Seed Proteins. Dordrecht: Kluwer.
Shewry PR, Jenkins J, Beaudoin F, and Mills ENC (2003) The classification, functions
and evolutionary relationships of plant proteins in relation to food allergens.
In: Mills ENC and Shewry PR (eds.) Plant Food Allergens. Oxford: Blackwell
Science.
 Protein Synthesis and Deposition
PR Shewry, Rothamsted Research, Harpenden, UK; University of Reading, Reading, UK
P Tosi, University of Reading, Reading, UK
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Seed storage proteins are synthesized on the endoplasmic
reticulum (ER) and cotranslationally transported into the
lumen.
Protein folding, assembly, and posttranslational modifica-
tions occur in compartments of the endomembrane system:
the ER, Golgi apparatus, and vacuole.
Proteins may be deposited in protein bodies derived from
the ER (many cereal prolamins) or vacuole (albumins,
globulins, and some cereal prolamins).
Seed storage protein genes are under complex regulation,
which ensures their tissue and temporal specificity of
expression but allows modulation related to the environ-
ment and nutrition.
Learning Objective
To understand the pathways and mechanisms of storage
protein synthesis, processing, trafficking, and deposition in
the endomembrane system of the seed.
Introduction
The protein content of seeds varies widely, from
10% of the
dry weight in cereals to
40% in soybean. The number of
individual protein components present is vast, but all seeds
contain well-defined groups of storage proteins, which account
for half or more of the total proteins. The storage proteins are
largely responsible for the nutritional quality and processing
properties of the grain and will, therefore, be the focus of this
article.
Four types of storage proteins occur widely in seeds, which
were initially defined based on their solubility properties and
molecular masses measured by sedimentation equilibrium
ultracentrifugation. The latter allows the calculation of sedi-
mentation coefficients (S20,w values) with higher values indi-
cating greater mass. Alcohol-soluble prolamins form the major
storage protein fraction in most cereal grain and two types of
globulins (7S and 11S), the major fractions in most dicotyle-
donous species, including legumes and oilseeds. However,
many of the latter also contain 2S albumin storage proteins.
Similarly, all cereals also contain storage globulins, with
proteins related to the 11S globulins of dicotyledonous plants
forming the major storage protein group in oats and rice.
The characteristics of these four types of proteins are summa-
rized in Table 1 and the trivial names used for different species
in Table 2. This article will focus on their synthesis and
deposition.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00106-6
Synthesis, Folding, Processing, and Deposition
of Seed Storage Protein
The vast majority of the proteins present in the mature cereal
grain, including all of the major groups of storage proteins, are
encoded by genes present on the chromosomes in the nucleus.
Smaller genomes are also present in two organelles, the mito-
chondrion and plastid, but these do not encode any major seed
proteins. Initiation of gene transcription is regulated by the
interactions between promoter elements located upstream of
the gene-coding regions and transcription factors, which are
proteins that interact with the promoter elements and with the
RNA polymerase enzyme that catalyzes the synthesis of mes-
senger RNA (mRNA) from the DNA template in the gene. The
mRNA is initially synthesized as a precursor form, which is
processed in the nucleus prior to being transported through the
nuclear membrane to the cytoplasm for translation.
Protein synthesis involves the translation of the nucleotide
sequence of the RNA (which is initially derived from that of the
genomic DNA) into the amino acid sequence of the protein. This
occurs on structures called ribosomes, which comprise com-
plexes of ribosomal RNA and proteins. Transfer RNAs bearing
amino acids and mRNAs are brought together on the ribosome
resulting in the formation of peptide bonds between adjacent
amino acids leading to the synthesis of the polypeptide chain.
Attachment of the mRNA to the ribosome occurs in the
cytoplasm and the translated protein may subsequently be
released into the cytoplasm. However, many proteins are syn-
thesized with an N-terminal signal peptide, which directs them
into the endomembrane system. This is because the translation
of the signal peptide causes the ribosome to attach to the outer
surface of the endoplasmic reticulum (ER), allowing the signal
peptide to direct the nascent (newly synthesized) protein
through the ER membrane into the lumen.
The major events in protein synthesis, and the way that the
sequence of nucleotides in the genomic DNA ultimately
determines the sequence of amino acids in the protein, are sum-
marized in Table 3. More detailed and highly readable accounts of
the various stages of protein synthesis are given in Buchanan et al.
Seed Storage Proteins Are Products of the
Endomembrane System
Seed storage proteins are present in the cells in dense protein
bodies. These originate from the endomembrane system of the
cell, which comprises the ER and structures derived from this,
including the Golgi apparatus and various types of vesicle,
which transport components to and from specific cellular
destinations. The parts of the endomembrane system that are
relevant to storage protein synthesis and deposition are shown
schematically in Figure 1, while the processing events are
summarized in Table 4. It should be noted that trafficking of
115
116 PROTEINS | Protein Synthesis and Deposition

Table 1 Summary of the characteristics of albumin, globulin, and prolamin storage proteins

2S albumins Soluble in water

Molecular mass (Mr) typically
1015 kDa
Processed posttranslationally to give large and small subunits
Two intrachain bonds within the large subunit and two interchain disulfide bonds
Not glycosylated
Specific components are rich in methionine
7S globulins Soluble in dilute salt solutions
Typically trimeric proteins of Mr 150190 kDa
Subunit Mr varies from
40 to 80 kDa but is typically
50 kDa
Subunits may be proteolytically processed and glycosylated
Contain little or no cysteine and methionine
11S Soluble in dilute salt solutions
globulins Typically hexameric proteins of Mr 300450 kDa. Subunits typically of Mr 60 kDa are posttranslationally processed to give Mr 40 kDa
(acidic) and Mr 20 kDa (basic) chains associated by one interchain disulfide bond
Low in cysteine and methionine
Prolamins Vary widely in structure, with subunit Mr ranging from
10 to 100 kDa
Include monomeric forms and high Mr polymers stabilized by interchain disulfide bonds
Presence of repeated sequences and regions (domains) rich in specific amino acids results in unusual amino acid compositions
Rich in proline and glutamine, poor in lysine and, in some cases, tryptophan, threonine, and methionine
No glycosylation or proteolytic processing
Soluble in alcohol/water mixture when native and/or reduced

Source: Shewry PR (2000) Seed proteins. In: Black M and Bewley JD (eds.) Seed Technology and Its Biological Basis, pp. 4284. Sheffield: Blackwell Science and Sheffield Academic
Press, with permission.

Table 2 Types of major storage proteins present in seed

crops and their trivial names Table 3 Summary of the major events in protein synthesis from
nuclear genes
Species Protein type Trivial name
Genomic DNA in the chromosomes of the nucleus consists of a linear
Cereals sequence of four nucleotides (bases). Individual genes comprise
Wheat (Triticum spp.) Prolamin Gliadin/ protein-coding sequences flanked by regulatory sequences. Coding
glutenin sequences specify the amino acid sequences of proteins in series of
Barley (Hordeum vulgare) Prolamin Hordein three bases called codons. The correspondence of codons to amino
Rye (Secale cereale) Prolamin Secalin acids is called the genetic code. Regulatory sequences control gene
Corn (Zea mays) Prolamin Zein expression by interactions with regulatory proteins (transcription
Sorghum (Sorghum bicolor) Prolamin Kafirin factors) and RNA polymerase enzymes
Oats (Avena sativa) Prolamin Avenin #
11S globulin Transcription of gene gives messenger (m)RNA with codon sequence
Rice (Oryza sativa) Prolamin reflecting that in the gene
11S globulin Glutelin #
Legumes mRNA is processed in the nucleus and transported via nuclear
Soybean (Glycine max) 7S globulin b-Conglycinin membrane to cytoplasm
11S globulin Glycinin #
Broad bean (Vicia faba)/pea (Pisum 7S globulin Vicilin mRNA binds to ribosome in the cytoplasm. Transfer RNAs bearing amino
sativum) 11S globulin Legumin acids recognize and bind to specific codons allowing the amino acids to
French bean (Phaseolus vulgaris) 7S globulin Phaseolin be linked in a specific sequence by peptide bonds. Thus the sequence
Peanut (Arachis hypogaea) 7S globulin Conarachin of codons determines the amino acid sequence of the protein
11S globulin Arachin #
2S albumin Translation of a signal peptide directs the ribosome to the endoplasmic
Oilseeds reticulum (ER) allowing translocation of the nascent protein into ER
Cottonseed (Gossypium hirsutum) 11S globulin Gossypin lumen
7S globulin Congossypin
2S albumin
Sunflower (Helianthus annuus) 11S globulin Helianthinin proteins through the endomembrane systems is a highly regu-
2S albumin lated process with the destinations of individual proteins being
Oilseed rape/canola (Brassica napus) 11S globulin Napin
determined by the presence of specific sequences or structural
2S albumin Cruciferin
features. In the absence of such signals, the protein is secreted
Source: Shewry PR (2003) Plant proteins. In: Thomas B, Murphy D, and Murray B from the cell by fusion of secretory vesicles derived from the
(eds.) Encyclopaedia of Applied Plant Science, pp. 889896. London: Elsevier Golgi apparatus with the plasma membrane. Hence, secretion
Science. is often referred to as the default destination.
 PROTEINS | Protein Synthesis and Deposition 117

Storage protein
synthesis occurs on
ribosomes attached to ER

RER
Smooth
ER

Transport Protein
in body
vesicles formation
within RER
Golgi
apparatus

Protein
body
formation

Secretory
vesicle Within
Plasma vacuole
membrane
Figure 1 Schematic summary of storage protein synthesis, processing, trafficking, and deposition in the endomembrane system of the cell.

Table 4 Processing of seed storage protein in the endomembrane

that is usually
20 amino acids) whose role is to lead the newly
system of the cell
synthesized (nascent) polypeptide through the ER membrane
ER Golgi apparatus Vacuole into the lumen, thus entering the endomembrane system. This
occurs by specific interaction of the signal sequence with a com-
2S albumins Folding, Proteolytic plex of proteins called the translocon. Once the nascent poly-
disulfide processing peptide emerges into the ER lumen, the signal sequence is
bond
removed by a specific enzyme (a signal peptidase) and the poly-
formation
peptide chain commences to fold into its three-dimensional
7S globulins Folding, N- Glycan Glycan
glycosylation, modification trimming, structure. These events, translocation, signal peptide cleavage,
formation of proteolytic and folding, occur when the protein is still undergoing synthesis
trimers processing and hence are termed cotranslational. Protein folding is almost
11S globulins Folding, Proteolytic certainly assisted by a complex of proteins and other factors, the
disulfide processing most well known of which is the molecular chaperone BiP
bond to give acidic (binding protein). In addition to assisting protein folding, BiP
formation, and basic also binds to malfolded proteins to prevent their escape from the
formation of chains, ER. Thus, it is often present in higher amounts in mutants in
intermediate formation of
which seed protein synthesis is affected (e.g., floury-2 and defective
trimers hexamers
endosperm B30 of maize) as well as during the period of max-
Prolamins Folding,
disulfide imum storage protein synthesis and deposition.
bond Most proteins synthesized on the ER also contain interchain
formation, and/or intrachain disulfide bonds. These bonds also form in
polymer the ER as an integral part of protein folding, and this may be
assembly assisted by the enzyme protein disulfide isomerase (PDI),
which catalyzes the formation, exchange, or reduction of disul-
fide bonds, depending on the substrate and conditions.
Synthesis, Folding, and Disulfide Bond Formation
Occur Simultaneously on the ER Storage Albumins and Globulins Are Deposited
in Storage Vacuoles
Seed storage protein is synthesized by mRNA associated with
polyribosomes present on the rough ER. The protein translated The 2S albumin and 7S and 11S globulin storage proteins are
from the mRNA differs from that deposited in protein bodies in transported via the ER lumen, Golgi apparatus, and vesicles to
the presence of a short N-terminal extension (the signal sequence specialized storage vacuoles where they form dense deposits.
118 PROTEINS | Protein Synthesis and Deposition
These vacuoles may subsequently divide to form protein bod-
ies. However, all three types of storage protein may also
undergo modification, either in the vacuole or during their
passage through the ER and Golgi apparatus.
The 2S albumins are synthesized as precursor proteins,
which undergo folding and disulfide bond formation in the
ER lumen. The precursors are then proteolytically processed in
the vacuole to generate the mature two-subunit structure. This
processing involves cleavage adjacent to specific asparagine
residues and the loss of one or more short peptide fragments
(from the protein N-terminus and C-terminus and between the
two subunits).
The 7S and 11S globulin subunit is also folded in the ER
lumen with the formation of a single intrachain disulfide bond
in the latter (7S globulins lack cysteine residues and hence have
no disulfide bonds). Both types of subunits are then assembled
into trimeric structures stabilized by noncovalent forces. There
has been considerable debate about the extent to which
globulin storage proteins are modified by the addition of
oligosaccharides (glycosylation). It is now established that
this rarely, if ever, occurs for 11S globulins but that some 7S
globulin subunits are modified by glycosylation of specific
asparagine residues (N-glycosylation). This occurs in several
stages, with the initial attachment of a high-mannose sugar
occurring cotranslationally (i.e., concurrent with folding and
disulfide bond formation). Subsequently, enzymes present in
the ER and Golgi apparatus may act to remove or add further
sugar residues to give a more complex structure with final
trimming occurring within the storage vacuole. Once within
the vacuole, the 11S globulins are proteolytically cleaved at an
internal asparagine residue to give acidic and basic chains that
remain associated by the single disulfide bond formed in the
ER. After this proteolytic processing, the intermediate dimers
formed in the ER are able to assemble to form the mature
hexameric structure.
The 7S globulin subunits may also be proteolytically pro-
cessed in the vacuole, but this depends on the species and the
precise protein subunit. For example, in bean (Phaseolus
vulgaris) and soybean (Glycine max), the 7S globulin subunits
do not undergo any proteolysis, while in pea (Pisum sativum),
some subunits are processed at one or two sites to give
polypeptide chains ranging in mass from
12 to 35 kDa, in
addition to unprocessed chains of mass
50 kDa. However,
the trimeric structure is not affected by these processing events.
Cereal Prolamins Are Deposited in Two
Types of Protein Body
Cereal prolamins do not undergo glycosylation or proteolytic
processing (except for cotranslational removal of the signal
sequence), but some subunits are assembled into high-
molecular-mass polymers stabilized by interchain disulfide
bonds. In wheat, these glutenin polymers may have masses of
between 1 and 10 MDa, and it is not known how such complex
multisubunit structures are assembled. However, it is probable
that assembly and interchain disulfide bond formation occur
rapidly after synthesis within the ER lumen. Subsequently, pro-
lamins are deposited in two discrete types of protein body.
In rice and maize (and probably also in sorghum and mil-
lets), prolamins accumulate directly within the lumen of the ER
to form a population of protein bodies surrounded by a mem-
brane of ER origin, which may remain studded with ribosomes.
In contrast, it appears that two separate pathways occur in wheat,
barley, and rye, with some prolamins being trafficked through
the ER and Golgi apparatus and deposited in storage vacuoles (as
described earlier for albumin and globulin storage proteins),
while others are retained in the ER as in maize and rice. It has
been suggested that the polymeric prolamins are preferentially
retained in the ER and monomeric forms transported to vacu-
oles, but it seems likely that the division is not so clear-cut. It is
also probable that the relative amounts of prolamins deposited
in the two types of protein body vary with the rate of protein
synthesis and with the developmental state of the tissue. Thus, it
can be envisaged that high levels of protein synthesis would lead
to a high level of accumulation within the ER lumen and that this
route would also be favored as the cells become distended with
deposits of starch and proteins. In wheat, the individual protein
bodies collapse during the later stages of seed maturation form-
ing a continuous matrix in the cells of the mature grain.
What Determines the Destination of Storage Proteins
in the Endomembrane System?
As discussed earlier, trafficking through the endomembrane
system is a highly regulated process with the destinations of
proteins being determined by their sequences and structures.
The retention of storage proteins within the ER or their
transport to the vacuole should, therefore, result from the pres-
ence of specific signals. A number of proteins are permanently
resident in the lumen of the ER, including PDI and BiP. In these
cases, retention results from the presence of a specific ER reten-
tion signal, which is a tetrapeptide located at the protein
C-terminus. The two most widespread tetrapeptides are KDEL
and HDEL, which correspond to the amino acids lysine or
histidine, aspartic acid, glutamic acid, and leucine. There is no
evidence that such a retention signal is present at the C-termini
of prolamins that are retained in the ER, and a different mech-
anism must therefore be sought. It has been suggested that BiP
plays a role in rice, by transiently binding to prolamins and
retaining them in the ER to allow assembly into protein bodies.
However, this mechanism has not so far been shown to be
applicable to other cereals. An alternative explanation is that
retention results from the insolubility of the proteins in aqueous
environments (i.e., the ER lumen) and their propensity to form
aggregates under such conditions. It can be envisaged that such
aggregates would accumulate directly in the ER lumen and then
be budded off to form protein bodies.
Some proteins are targeted to the vacuole by the presence of
specific prosequences that are removed by posttranslational
proteolysis once the protein arrives in the vacuole. These pro-
sequences may be at the protein N-terminus, as in the barley
cysteine proteinase aleurain and the sweet potato storage pro-
tein sporamin (which is stored in tubers) or at the C-terminus
as in lectins; more rarely, a noncontiguous series of residues
form a tridimensional structure in the mature protein, which
acts as a vacuolar sorting signal. None of the 2S albumins, 7S
globulins, 11S globulins, or prolamins that are deposited in
 PROTEINS | Protein Synthesis and Deposition 119

vacuolar protein bodies have cleavable prosequences that are

involved in vacuolar targeting. However, it has been proposed
that aggregation of 7S and 11S globulins into electron-dense
aggregates in the Golgi is a prerequisite for ensuring that they
are sorted into dense vesicles and ultimately deposited into the
vacuole, whereas other proteins are sorted into different types
of vesicles with different destinations. This sorting may require
the presence of specific hydrophobic residues, which mediate
the proteinprotein interactions. However, the precise mecha-
nisms that determine the final destination of seed storage pro-
teins, whether to ER-derived or vacuolar protein bodies, are
still poorly understood.

Organization of Storage Proteins in Protein Bodies

(a)
The 7S and 11S globulins have similar three-dimensional struc-
tures with the 11S hexamers essentially corresponding to
dimers of the trimeric 7S proteins. This similarity presumably
facilitates their efficient packaging in the same protein bodies.
However, protein bodies may also contain separate phases, or
inclusions, containing specific proteins or other components.
For example, the protein bodies of castor bean consist of a matrix
of 2S albumins and other proteins (including lectins) with crys-
talline inclusions of 11S globulins and noncrystalline globoid
inclusions consisting of phytin (calcium and magnesium salts of
myoinositol hexaphosphoric acid). Phytin globoids are also pre-
sent in protein bodies of other dicotyledonous seeds and in
aleurone cell protein bodies of cereals. The different routes,
taken by prolamins and glutelins (11S globulin homologues)
(b)
in rice lead to the formation of two separate populations of
protein bodies called PB-I (ER-derived containing prolamins)
and PB-II (vacuolar-derived containing glutelins). However, in
wheat and oats, these two types of protein are located in the same
protein body, with prolamin inclusions in a globulin matrix in
oats and vice versa in wheat (Figure 2). This segregation has been
suggested to result in oats from the fusion of ER-derived protein
bodies containing prolamins with vacuolar protein bodies con-
taining globulins.
It has also been shown that different types of prolamin
(zein) are concentrated in the outer and inner parts of protein
bodies of maize, although in this case, the location appears to
(c)
result from the pattern of development of the bodies in the
endosperm cells. Figure 2 Protein bodies in developing starchy endosperm cells of
cereals. (a) Rice at 7 days after flowering, showing two populations of
protein bodies. PB-Is are spherical vesicles delimited by a single unit
Assembly of Glutenin Polymers in Wheat membrane derived from the ER and contain prolamins. PB-IIs are
amorphous, derived from vacuolar deposition and contain globulins/
Polymer assembly is of particular interest in wheat as the glutelins. Reproduced from Yamagata H and Tanaka K (1986). The site of
formation of high-molecular-mass glutenin polymers under- synthesis and accumulation of rice storage proteins. Plant and Cell
pins the processing properties of flour, by conferring elastic Physiology 27: 135145, with permission. (b) Oats at 8 days after anthesis
properties to dough. These polymers comprise of two major showing light-staining deposits of prolamin (labeled with 10 nm colloidal
types of subunit, called high-molecular-weight (HMW) and gold) as inclusions in protein bodies containing globulins
low-molecular-weight (LMW) subunits. These subunits are (5 nm colloidal gold). Reproduced from Lending CR, Chesnut RS, Shaw KL,
and Larkins BA (1989). Immunolocalization of avenin and globulin
linked by interchain disulfide bonds that are presumably ini-
storage proteins in developing endosperm of Avena sativa L. Planta 178:
tially formed during folding in the ER (as discussed earlier). 315324, with permission. (c) Wheat at 11 days after flowering showing
The complexity and size of the glutenin polymers have limited inclusions of triticin (dark staining, labeled I) within a matrix of prolamin
attempts to determine their precise structures, but current evi- (M). The bars are 1.0 mm in (a) and 0.5 mm in (b) and (c). Reproduced from
dence is consistent with a model in which the HMW subunits Bechtel D, Wilson JD, and Shewry PR (1991). Immunocytochemical
form the backbone, linked mainly head to tail, while the localization of the wheat storage protein triticin in developing endosperm
LMW subunits form branches. In addition, glutenin polymers tissue. Cereal Chemistry 68: 573577, with permission.
120 PROTEINS | Protein Synthesis and Deposition
interact strongly with each other and with the monomeric
gliadins by noncovalent forces, particularly hydrogen bonds.
This combination of covalent and noncovalent interactions,
together with the intrinsic structures of the individual subunits,
results in the viscoelastic properties associated with wheat
dough and gluten.
The limited information that is available indicates that
glutenin polymers have defined structures, with specific inter-
actions between individual subunits. Similarly, it is clear that
some allelic forms of HMW subunits are associated with the
formation of larger polymers, leading to positive effects of
quality. However, the mechanisms responsible for defining
specific structures are not known.
Finally, the proportion of high-molecular-mass glutenin
polymers increases dramatically during the desiccation phase of
grain development, suggesting that polymer formation is also
driven by dehydration. This is consistent with the model that
glutenin polymers are stabilized by strong arrays of hydrogen
bonds, but rearrangements of disulfide bonds may also occur.
Regulation of Storage Protein Synthesis
Tissue Specificity
Storage protein synthesis in seeds is strictly regulated with
respect to timing, tissue specificity, and, to a lesser extent, nutri-
ent availability. In addition to the embryo, which is a diploid
zygotic tissue that gives rise to the seedling, all seeds also contain
an endosperm, which is triploid and is derived from a second
fertilization event. However, the size of the endosperm varies
greatly, forming 8090% of the mature cereal grain but being
largely broken down during development in soybean and many
other seeds. Similarly, whereas some seeds such as quinoa and
amaranth contain a perisperm (derived from the maternal
nucellar tissue), this fails to develop and is absorbed in most
species. In all seeds, at least one of these three tissues functions
as a storage tissue, and in some cases, more than one. The central
part of the endosperm is the major storage tissue in cereals,
accumulating protein (prolamins with varying amounts of
11S-type globulins), and starch. However, 7S globulins and oil
are also stored in the outer cells of the endosperm (the aleurone
layer) and the single cotyledon (scutellum) of the embryo. The
cotyledons of the embryo form the major storage tissue in grain
legumes (including soybean, peanut, lupin, and pea),
sunflower, and oilseed rape/canola, while proteins and oil are
stored in the endosperm and embryo of quinoa with starch
being stored in the perisperm.
It should also be noted that the synthesis of seed storage
proteins has never been detected in nonseed tissues of the
plant, whether by determination of gene expression (i.e.,
mRNA populations) or by protein accumulation.
Temporal Regulation
Seed development proceeds through a series of genetically
programmed stages, starting with cell division and differentia-
tion to establish the basic structure of the component organs
and tissues (i.e., the embryo and endosperm). The synthesis
and accumulation of storage products, including proteins,
occur during the subsequent expansion of the cells of the
storage tissue(s) and cease when these cells mature and desic-
cate. The onset and duration of the phases vary widely between
species and genotypes and are greatly affected by the environ-
mental conditions. In small grain cereals (wheat and barley)
grown in the United Kingdom, storage protein synthesis is
usually first detected in appreciable amounts at
1214 days
after pollination and is most active over the next 23 weeks.
However, synthesis may be initiated as early as 6 days in plants
grown in hot and dry climates. In maize (corn) grown in North
America, storage protein synthesis occurs between
3 and 7
weeks after pollination.
Storage protein fractions are, of course, not single proteins
but mixtures of components, which may be encoded by several
multigene families. In general, these genes show highly coor-
dinated patterns of expression, with only small differences in
their timing and relative levels of expression during develop-
ment. For example, in sunflower, the expression of genes for
11S globulin storage proteins (helianthinins) is detected
slightly before that of genes for 2S albumins, but in wheat
and barley all groups of prolamin storage proteins show sim-
ilar expression patterns.
Environmental Control
Environment has a major impact on seed development, includ-
ing storage protein synthesis, with higher temperatures
generally increasing the rate of development. In addition,
specific heat-shock effects may occur when temperatures
exceed
35  C: these can result in detrimental effects on the
quality of wheat grown in Australia and some other parts of the
world. The most dramatic effect of environment on storage
protein synthesis is that of nutrition. It is well established
that the protein content of seeds is regulated by the availability
of nitrogen, with storage protein acting as a sink for nitrogen,
which is not required by the plant for other purposes. For
example, increasing the total nitrogen content of barley grain
from
1% to 3% is associated with an increase in prolamin
(hordein) storage proteins from
35% to 50% of the total
grain nitrogen. This effect can be exploited to grow grain with
defined protein contents for specific end uses (e.g., low protein
for malting barley and high protein for bread-making wheat).
However, almost all storage proteins also contain sulfur in
the form of the amino acids cysteine and methionine. Many
species are able to adjust to variation in the relative availabil-
ities of nitrogen and sulfur by having specific types of seed
storage proteins with high and low contents of sulfur amino
acids. Thus, the 2S albumin storage proteins of oilseed rape
and sunflower have higher contents of cysteine and methio-
nine than the globulin storage proteins, and the synthesis of
the former is therefore reduced under conditions of low sulfur
availability (Figure 3(a) and 3(b)). Similarly, restricted avail-
ability of sulfur favors the synthesis of the sulfur-poor C hor-
deins of barley (Figure 3(c)) and o-gliadins of wheat over
other more sulfur-rich prolamins, which can have adverse
consequences for grain quality in the latter species.
Regulation of Storage Protein Gene Expression
The regulation of seed storage protein genes usually occurs at
the level of gene transcription, although some fine-tuning can

1 been identified, the precise mechanisms of storage protein
2
3
(a) + (b) + (c) + in food products. Furthermore, it is becoming clear that the
Figure 3 The effect of sulfur on the total seed proteins of oilseed rape
(a) and sunflower (b) and on the prolamin (hordein) proteins of barley
(c). Note the presence of S-rich low-molecular-mass proteins (probably
2S albumins) in oilseed rape and sunflower (bracket) and of S-poor C
hordeins (2) in barley. Parts (a) and (b) are reproduced from Spencer D,
Rerie WG, Randall PJ, and Higgins TJV (1990). The regulation of pea
seed storage protein genes by sulfur. Australian Journal of Plant
Physiology 17: 355363, with permission. Part (c) is reproduced from
Shewry PR, Franklin J, Parmar S, Smith SJ, and Miflin BJ (1983) The
effects of sulfur starvation on the amino acid and protein composition of
barley grain. Journal of Cereal Science 1: 2131, with permission.
occur at the level of translation of the mRNA into proteins.
Consequently, interest has focused on the identification of
regulatory sequences present upstream of the gene sequences
that encode storage proteins (called 50 upstream sequences)
and of specific proteins that bind to these (transcription fac-
tors). This has led to the identification of short conserved
nucleotide sequences (often called boxes) that regulate expres-
sion of the genes.
Comparison of the 50 upstream sequences of a number of
genes encoding 11S storage globulins has led to the identifica-
tion of a 28 base pair (bp) legumin box. Deletion of a central
7-nucleotide motif from the center of this box resulted in a
drastic reduction in the gene expression in transgenic plants,
demonstrating its importance. Related boxes are also present
in the 50 upstream sequences of 7S globulin and 2S albumin
genes. However, further studies have demonstrated the pres-
ence of several additional upstream regulatory sequences that
may modulate storage globulin gene expression.
Similar studies have shown that many prolamin genes also
contain a conserved sequence of
30 bp located about 300 bp
upstream of the gene-coding region. This is called the
endosperm box or prolamin box and is unrelated in
sequence to the legumin box. The prolamin box actually
comprises two conserved sequences separated by a more vari-
able region. One of these conserved sequences is thought to be
responsible for conferring endosperm specificity, while the
other may play a role in the regulation of the gene. However,
additional regulatory sequences are undoubtedly also present
including a short sequence (the E motif) in the a-zein genes of
PROTEINS | Protein Synthesis and Deposition 121
maize, which binds the protein encoded by the Opaque 2 locus.
Mutation of this locus results in reduced a-zein synthesis and a
high lysine phenotype. Although several other proteins that
bind to regulatory sequences of storage protein genes have also
gene regulation remain poorly understood.
Conclusions
It is clear that we still have much to learn about the precise
mechanisms of storage protein synthesis, trafficking, and depo-
sition, although the broad principles are now understood.
A detailed understanding of these mechanisms is important
as interactions (i.e., between proteins and with other grain
components) established during synthesis and deposition
may ultimately affect the functional properties of the proteins
synthesis, assembly, deposition, and interactions of seed pro-
teins may be affected by environmental factors that ultimately
impact on seed quality. Identifying the major sites of these
effects and the critical stages of seed development should
allow them to be predicted and ultimately manipulated to
give a higher level of stability.
It is also important to understand the mechanisms of seed
protein synthesis, trafficking, and deposition in order to facil-
itate the application of genetic engineering to improve grain
quality. It is essential that any novel or mutant proteins
expressed in transgenic plants should be accepted by the
secretory system as authentic and transported to their correct
destination. Any errors in folding or failure to contain the
correct targeting information could lead to protein turnover
or accumulation in incorrect compartments (e.g., ER instead of
storage vacuoles in dicotyledonous seeds) with adverse effects
on grain development, quality, and yield.
Exercises for Revision
Summarize the compartments of the endomembrane sys-
tem of the plant cell in relation to the trafficking and
processing of seed storage protein.
Discuss the origins of protein bodies in cereal and dicoty-
ledonous seeds.
Discuss the pathways and mechanisms of processing and
deposition of different types of storage proteins: albumins,
globulins, and prolamins.
Exercises for Readers to Explore the Topic Further
Identify the structure/sequences that determine storage pro-
tein targeting and trafficking in the endomembrane system.
Determine the organization and interactions of storage pro-
teins within protein bodies.
Determine the precise mechanism of storage protein gene
regulation in the context of grain development, the nutri-
tional status of the plant, and abiotic stress.
122 PROTEINS | Protein Synthesis and Deposition
See also: Proteins: The Protein Chemistry of Cereal Grains; The
Protein Chemistry of Dicotyledonous Grains; Wheat-Based Foods:
Cereals: Overview of Uses: Accent on Wheat Grain.
Further Reading
Bechtel D, Wilson JD, and Shewry PR (1991) Immunocytochemical localization of the
wheat storage protein triticin in developing endosperm tissue. Cereal Chemistry
68: 573577.
Bewley JD and Black M (1995) Seeds Physiology of Development and Germination, 2nd
edn. New York: Plenum Press.
Buchanan BB, Gruissem W, and Jones RL (2000) Biochemistry and Molecular Biology
of Plants. Rockville, MD: American Society of Plant Physiologists.
Larkins BA and Vasil IK (1997) Cellular and Molecular Biology of Plant Seed
Development. Dordrecht: Kluwer Academic.
Lending CR, Chesnut RS, Shaw KL, and Larkins BA (1989) Immunolocalization of
avenin and globulin storage proteins in developing endosperm of Avena sativa L.
Planta 178: 315324.
Muntz K and Shutov AD (2002) Legumains and their functions in plants. Trends in Plant
Science 7: 340344.
Shewry PR (2000) Seed proteins. In: Black M and Bewley JD (eds.) Seed Technology
and its Biological Basis, pp. 4284. Sheffield: Blackwell Science and Sheffield
Academic Press.
Shewry PR (2003) Plant proteins. In: Thomas B, Murphy D, and Murray B (eds.)
Encyclopaedia of Applied Plant Science, pp. 889896. London: Elsevier
Science.
Shewry PR and Casey R (1999) Seed Proteins. Dordrecht: Kluwer Academic.
Shewry PR and Halford NG (2002) Cereal seed storage proteins: Structures, properties
and role in grain utilization. Journal of Experimental Botany 53: 947958.
Shewry PR, Franklin J, Parmar S, Smith SJ, and Miflin BJ (1983) The effects of sulphur
starvation on the amino acid and protein compositions of barley grain. Journal of
Cereal Science 1: 2131.
Shewry PR, Halford NG, Tatham AS, Popineau Y, Lafiandra DL, and Belton PS (2003)
The high molecular weight subunits of glutenin and their role in determining
wheat processing properties. Advances in Food and Nutrition Research
45: 221302.
Spencer D, Rerie WG, Randall PJ, and Higgins TJV (1990) The regulation of pea seed
storage protein genes by sulphur. Australian Journal of Plant Physiology
17: 355363.
Tosi P (2012) Trafficking and deposition of prolamins in wheat. Journal of Cereal
Science 56: 8190.
Yamagata H and Tanaka K (1986) The site of synthesis and accumulation of rice storage
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 Nitrogen Metabolism
CA Atkins, The University of Western Australia, Crawley, WA, Australia
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 287292, 2004, Elsevier Ltd.
Topic Highlights
Mature plant seeds contain nitrogen-based nutrient
reserves, particularly proteins.
Nitrogen is transported into grain or seed, where it is
metabolized into amino acids.
Most nitrogen entering the seed is sued to synthesize stor-
age proteins, which are utilized by the germinating seedling
(or enter into food processing).
Learning Objectives
To understand the dynamics of nitrogen metabolism that
results in the storage of useful plant proteins in the mature
grain.
To understand the transportation of nitrogen-containing
solutes into the seed, in relation to the concept of sink
strength.
To understand the metabolism of nitrogen within the seed
and the formation of storage reserves.
Introduction
The success that flowering plants enjoy is in no small way a
consequence of the effectiveness of their seed as a perennating
organ. The embryo is in a metabolically dormant state and is
accompanied by a reserve of nutrients that can be mobilized on
germination to support the activated meristems of the new
plant. The diversity of composition exhibited by mature seeds
reflects variation in the size and nature of this stored reserve.
Some are high in carbohydrate, others in protein, and others in
oil. Yet all are viable and are able to convert the reserves into
translocatable solutes with a balance of C and N that supports
the active root and shoot meristems before uptake from soil
provides an alternative source of nutrients and before the
seedling is autotrophic for C through photosynthesis. The
reserves are fashioned from organic assimilates that supply C,
N, and S through the long distance translocation channels of
the vegetative structure of the plant.
Metabolism of N has been studied in a restricted range of
seed types, mostly those of cereals, legumes, and oilseeds of
agricultural importance and more recently in the model species
Arabidopsis thaliana. Because of the high proportion of protein
stored in legume seeds these have received most attention in
this regard. However it is well to realize that plants produce an
almost bewildering array of N-containing compounds, many
of which are amino compounds that accumulate in seeds and
are likely to impart unique nutritional, toxicological and phar-
macological properties. Our understanding of this whole area
in angiosperms is thus limited and fragmentary, and at this
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00104-2
stage we must infer the commonality of processes that are so
far only adequately documented for a few species.
Nitrogen Nutrition of Developing Seeds
Before considering seed nutrition it is useful to review the basic
structural features of seeds and the consequences of this struc-
ture to solute import into and transfer within the component
tissues of a seed as it develops. The outer seedcoat tissues are
maternal while the endosperm and embryo within this coat are
filial. There is no vascular connection between the two. Thus,
whether stored reserves of the seed accumulate in the endo-
sperm, the embryo, or associated tissues, in all cases solutes
that supply C and N (assimilates) pass across an apoplastic
compartment. It follows that in receiving translocated solutes
before they pass to the embryo the seedcoat may regulate both
the rate and nature of solute transfer.
Although developing seeds and their surrounding support-
ing structures may have the capacity for photosynthesis during
early stages of development, this is limited. Consequently their
major source of C is assimilates translocated in phloem and
xylem. These are principally in phloem as sucrose, longer
raffinose-based oligosaccharides, or in some species cyclitols,
like mannitol and sorbitol. The same translocation streams
supply N but typically phloem contains 1020 times more N
than xylem. However, even though both streams may carry a
wide array of amino acids, as well as other N-solutes (e.g.,
proteins, nucleic acids, plant growth regulators), usually one
or two low molecular weight organic compounds predomi-
nate. For many dicotyledonous species, including temperate
legumes, these are the amides, glutamine and asparagine or
their parent acids, glutamate, and aspartate, especially in
cereals. In some species arginine, proline, or non-protein
amino compounds like homoserine, ornithine, citrulline,
canavanine, and putrescine predominate. In others, particu-
larly nodulated legumes of tropical origin, the ureides, allan-
toin, and allantoic acid, account for almost all the N in xylem
and a significant component in phloem-N. There are some very
unusual N-solutes that translocate the bulk of N, for example
djenkolic acid (two cysteines bridged at their S groups by a
methylene) occurs at very significant levels in the xylem of
some Acacia spp., and no doubt the plant kingdom employs
many other forms of translocated N that have yet to be discov-
ered. Thus protein synthesis in sink organs does not receive a
mixture of amino acids that can be simply used to form pro-
tein, but rather a mixture in which just one or two solutes
provide the bulk of N that has to be metabolized to generate
the appropriate proportions of the 20 protein amino acids. In
their passage from sites of phloem unloading to the embryo,
N-solutes may be transiently stored, for example in the
123
124 PROTEINS | Nitrogen Metabolism
seedcoat, and are metabolized in highly specific ways within
the tissue compartments they traverse.
There seems little doubt that almost all the N entering seeds is
in an organic form. Although the literature records quite high
levels of ammonium ion in both xylem and phloem, it is not
likely that this is the norm and may, in many cases, be a conse-
quence of hydrolysis of the amide groups of glutamine and
asparagine during or subsequent to collection of vascular exu-
dates. Xylem transports considerable quantities of nitrate from
the root system to the major transpiring surfaces of the shoot but
negligible quantities of this inorganic ion are transferred to
phloem. As a result, fruit tissues and especially developing seeds
are unlikely to receive significant levels of nitrate and conse-
quently exhibit low or negligible capacity for its assimilation.
Early in legume fruit/seed development when a significant
proportion of translocated N is delivered in xylem, the C:N
ratio of incoming solutes is more narrow than that required by
the developing tissues, and excess N is transiently stored. The
endosperm in many dicotyledonous seeds accumulates a sig-
nificant proportion of this N as ammonium ion and levels
approaching 0.1 M have been recorded for periods of 34
weeks in white lupin (Lupinus albus) seeds. Although the
ammonium is promptly reassimilated when the developing
embryo expands to fill the endospermic apoplast, such high
levels are normally regarded as extremely toxic and mecha-
nisms that prevent this ammonium entering the embryo to
any great extent must be functioning at this time. In pea
(Pisum sativum), the major forms of accumulated N in the
endosperm are homoserine, glutamine, and alanine, in cow-
pea (Vigna unguiculata) endosperm there is negligible ammo-
nium ion but 3055% of the N is histidine, while in chickpea
(Cicer arietinum) ammonium and citrulline predominate. The
significance of ammonium or other N solutes that accumulate
to such high levels in endosperm when embryo development is
characterized by cell division is unknown, but it is interesting
to note that coconut endosperm (milk), used in pioneering
studies of plant cell and tissue culture, contains high concen-
trations of N, including ammonium ion, along with plant
growth regulators.
A positive correlation has been established between rates of
ammonia emission (i.e., N loss) and ammonium concentra-
tion in leaves. There are no emission data available for devel-
oping seeds/fruits, but given the high levels of ammonium ion
that may accumulate in liquid endosperm it seems possible
that transient periods of N loss occur due to its volatilization. It
is perhaps not too surprising then that the highest rates of
ammonia emission in barley occur during grain filling.
The mix of solutes in phloem is not constant during repro-
ductive growth, the C:N ratio changing to more closely match
the C and N requirements of the developing seed. In white
lupin, sucrose levels in phloem decline progressively after
anthesis while the concentration of amino compounds
increases, almost doubling over the period of grain filling.
This pattern of change is consistent with a declining rate of
photosynthesis and increasing senescence of leaves, with
release of protein-N as amino compounds, during reproduc-
tive development. However, further studies with lupin have
shown that in fact the phloem N level also increases to narrow
the C:N ratio as a consequence of solute specific mechanisms
that achieve an enrichment of phloem with asparagine as the
translocation stream passes through stem segments immedi-
ately before entering the inflorescence. Similar specific
exchanges of amino acids have also been found for developing
cowpea fruits/seeds. The precise tissue or cellular site where
these transfer mechanisms act and the molecular basis for their
regulation is unknown but a role for transfer cells has been
inferred.
The developing seed is supported on or within fruiting
structures that not only bear the vasculature which brings the
translocation streams to the seed but which serve also as inter-
mediate sites for metabolism and transient accumulation of
nutrients, including N. For example, in the period from anthe-
sis to 11 days after anthesis, in cowpea fruits 21 mg N enters in
phloem and 17 mg N in xylem. Of this N less than half
(15 mg) reaches the developing embryo and endosperm, the
rest is sequestered into the pod wall (16 mg) and the testa
(7 mg). Importantly, however, the incoming N-solutes are
metabolized extensively before being transferred to the
embryo. At this early stage of development xylem-N is
70%
ureide and phloem-N
20% ureide, but ureides account for
less than 20% in the podwall and testa and no more than 2%
of soluble-N in the liquid endosperm or embryo. Later in
development (1222 days after anthesis) phloem accounts
for 80% of N entering the fruit and, although the ureide level
is low, ureide-N accumulates preferentially in the seedcoat.
Despite this enrichment literally no ureide is transferred to
the apoplast surrounding the embryo. Rather, ureide-N is con-
verted to glutamine and this is the major source of N for
protein synthesis. Similar observations have been made for
soybean, another plant that translocates significant levels of
ureide-N in xylem and to a lesser extent in phloem.
In pea a major N-solute entering the seedcoat in phloem is
homoserine and, although this may accumulate transiently in
endosperm, most is converted to threonine and glutamine in
the seedcoat during embryo expansion. Even in species that
have an asparagine-rich phloem stream reaching the seedcoats,
a significant proportion of asparagine-N is converted to
glutamine-N during passage to the apoplast and embryo.
Whether this apparent preference for N as glutamine in the
embryos nutrition reflects some metabolic requirement or the
specificity of transporters involved in transferring solutes
across tissue compartments of the seed has yet to be clearly
resolved. In maize the endosperm of the developing kernel
supplied assimilates through the pedicel and even though the
major phloem N-solute is aspartate this is largely metabolized
with its N being transferred to glutamine. The maize pedicel
expresses a unique cytosolic form of glutamine synthetase,
GSp1, which is developmentally regulated, increasing as storage
protein synthesis increases. Glutamine is the major amino acid
released from the pedicel to the apoplast surrounding the
endosperm and a transporter expressed in the basal endosperm
transfer cells apparently enhances its uptake. Thus specificity of
both metabolism and transport combine to ensure that a pre-
ferred N-solute reaches the embryo tissues.
Transport of N-Solutes within Seeds
Rates at which solutes that supply C and N to the embryo are
transported are likely to contribute to determining the

potential accumulation of assimilates or sink strength of seeds
and consequently grain yield and harvest index. Studies with
four cultivars of bean (Phaseolus vulgaris) that differed in their
rates of seed growth found that although sink size, as assessed
by surface area and volume of cotyledons, were major deter-
minates of growth rate, the rate of transfer of dry matter was
also a contributing factor. Not surprisingly, maximal sucrose
fluxes into cotyledons were positively correlated with levels of
expression of the sucrose/H symporter and accompanying
H-ATPase in the dermal cells facing the apoplast between
the cotyledons and the seedcoat. Specifically in relation to
N nutrition, activity of amino acid transporters has been sug-
gested to regulate the rate of seed protein synthesis. Although
most research into solute unloading from phloem and trans-
port within developing seeds has involved sucrose, some of the
amino acid transporters expressed in seed tissues have been
identified and their location studied in relation to likely
function.
In pea, two cDNA clones (PsAAP1 and 2) belonging to
the AAP (amino acid permease) family of H/amino acid
co-transporters have been isolated from cotyledons. Like the
sucrose transporter, PsAAP1 is highly expressed in special-
ized epidermal transfer cells abutting the seedcoat and
shows broad specificity, mediating transport of acidic, neu-
tral, and basic amino acids. Similar broad-spectrum mem-
bers of the AAP family have been described for developing
Arabidopsis and castor bean seeds. In Arabidopsis AtAAP1 is
localized to endosperm and cotyledons while AtAAP2 is
restricted to the vasculature of siliques and the funiculus.
Thus, AtAAP2 is thought to mediate exchange of amino
acids from xylem to phloem. Such an exchange has been
identified as a specific asparagine transfer in the upper stem
segments and raceme of lupin to enrich phloem with N,
narrowing the C:N ratio to match more closely seed
demands for these commodities.
The broad substrate specificity found for most plant amino
acid transporters does not conflict with the idea that translo-
cated N is characterized by just one or two amino acids. Most
of the amino acid transporters from plants have been charac-
terized by yeast mutant complementation or functional analy-
sis in Xenopus oocyte assays and specificity in vivo may be quite
different. Indeed, differences in affinity for amino acid species
have been described. Furthermore, specific transporters are
likely to be found when more extensive genomic data is avail-
able for a wider range of plants.
As noted above seed tissues produce and may transiently
accumulate significant amounts of ammonium ion. It appears
that this is a feature of their metabolism throughout devel-
opment, raising the question of transport mechanisms for
the ion within seed tissues. Preliminary data indicates that
genes homologous to the high affinity ammonium trans-
porter family (AMT), AtAMT1 from Arabidopsis, are not
expressed in the seedcoat and cotyledons of developing
lupin seeds. However, the recently described high affinity
AMT2 transporter gene is expressed in Arabidopsis siliques
and this may be the transporter that functions in seeds. In
view of the widely diverse levels of ammonium that form in
seed tissues at different stages of development it seems likely
that other transporters/channels with quite different kinetic
features are also involved.
PROTEINS | Nitrogen Metabolism 125
The Arabidopsis genome has revealed transporters for other
types of N-solute that are expressed and could have significant
roles in seed metabolism. These include oligopeptide trans-
porters, such as AtPTR2-B, that might be responsible for the
movement of peptides released from storage proteins during
germination to new meristems or in the transport of signal
peptides from sites of synthesis to sites of action. The possible
importance of peptide transport in seed development was
highlighted in a study where antisense lines for AtPTR2-B
were generated in Arabidopsis. The transgenic plants showed
normal silique development but formed fewer, larger seeds
than controls. Specifically about half of the embryos were
arrested at an early stage of their development, suggesting
peptide transport was essential for N nutrition at this time or
that transfer of some bioactive peptide, essential for embryo-
genesis, was limiting.
Plants also express high affinity transporters specific for
purines and purine derivatives (PUPs) including bases, nucle-
osides, purine alkaloids, and cytokinins. Northern analysis for
AtPUP1 in Arabidopsis has however shown only a low level of
expression in developing siliques compared to other tissues
and their functional significance in seed development or in
the mobilization of N during germination remains to be
established.
A novel superfamily of transporters has recently been
described for allantoin and other oxidized derivatives of het-
erocyclic N solutes in Arabidopsis. In view of the predomi-
nance of ureide-N in translocation streams serving developing
fruits in certain tropical legume species, but with the meta-
bolic requirement that ureide-N is not transferred from the
maternal tissue to the embryo, it will be interesting to see
whether or not fruit and seed tissues express these trans-
porters. Enzymes of purine oxidation (urate oxidase and
allantoinase) have been demonstrated at elevated levels dur-
ing germination of a number of seeds, and it has been
inferred that purines are released due to intense nucleic acid
hydrolysis. If indeed allantoin provides a supplementary
source of translocated N to the newly formed root and
shoot apices then the novel allantoin transporters may be
important in phloem loading at this time.
Primary N Metabolism
The primary pathways of N metabolism in seeds are the reac-
tions that utilize incoming N-solutes to form the 20 protein
amino acids. As noted above N-solutes translocated to tissues
surrounding a developing seed or to the embryo itself are
characterized by one or two predominant compounds.
Among these asparagine and glutamine appear to be the
most common. Although some incoming asparagine is incor-
porated directly into seed protein the majority is metabolized
(in white lupin more than 80% over 13 weeks of develop-
ment). Supplying 15N (amide)-labeled asparagine to develop-
ing lupin fruits in phloem resulted in label being recovered in
15 amino acids following hydrolysis of seed protein. In addi-
tion to asparagine, glutamine, aspartate, and glutamate, those
labeled included arginine and histidine, those derived from
pyruvate (alanine, leucine and valine) and 3-phosphoglycerate
(glycine and serine), those of the aspartate family (lysine and
126 PROTEINS | Nitrogen Metabolism
isoleucine) as well as aromatic amino acids (tyrosine and
phenylalanine). Clearly seeds express all the major pathways
for synthesis of protein amino acids.
There are two potential pathways releasing asparagine-N to
amino acid synthesis. Hydrolysis of the amide-group by aspar-
aginase produces ammonia and aspartate, the latter providing
an amino donor to the many aminotransferases (AT) that have
been detected in seeds. The second pathway utilizes an
asparagine-AT that transfers the amino group to glyoxylate
forming glycine and 2-oxosuccinamic acid. The acid is reduced
to 2-hydroxysuccinamate and deamidated to ammonia and
malate. Although both potential pathways are expressed in
developing pea fruits, asparagine-AT is largely confined to
pod tissue and is very low in developing seeds. Asparaginase
on the other hand is expressed at high levels in seedcoats early
in development, transiently in endosperm and in filling coty-
ledons of legume seeds. The same could be said for predomi-
nant non-amino acid solutes like the ureides. Although there
are a number of enzymic routes for the degradation of allan-
toin that are expressed in different plants, differing even
between cultivars, the N is released as ammonia. Thus reassi-
milation of ammonia is a major activity of seed tissues whether
in the seedcoat, endosperm, or the components of the embryo.
Not surprisingly, both glutamine synthetase (GS) and gluta-
mate synthase (GOGAT) activities are expressed in seed tissues,
providing a high affinity utilization mechanism for ammonia
that is likely to function in reassimilation throughout seed
development. Seeds express glutamate dehydrogenase (GDH)
activity and this may also participate in reassimilation. While
the role of GDH has been the subject of considerable debate
and conflicting opinion the consensus view that has emerged
for most tissues appears to be one of oxidation of glutamate
rather than synthesis. However, the very high concentrations of
ammonium ion that form transiently in the endosperm of
some seeds would be kinetically favorable for GDH to func-
tion, for a short while at least, in its assimilation.
Among the protein amino acids that are synthesized largely
in situ to meet the needs of seed storage protein accumulation
are those of the aspartate family; namely lysine, threonine,
isoleucine, and methionine. Not surprisingly expression of
the first committed step in the aspartate pathway, catalyzed
by a bifunctional aspartate kinase (AK)-homoserine dehydro-
genase (HSDH) in Arabidopsis, is coincident with storage pro-
tein synthesis in the embryo. Cereals are deficient in lysine and
in some, threonine and tryptophan, while legume seeds are
poor sources of S-amino acids, particularly methionine. As a
consequence there has been considerable research into regula-
tion of the pathway for these essential amino acids. There
have also been numerous attempts to exploit this knowledge
to enhance the nutritional and/or processing qualities of both
cereal and pulse grains. AK activity is subject to feedback inhi-
bition by pathway products, specifically lysine or threonine.
However, in the high lysine opaque-2 mutants of maize very
significant levels of the aspartate family accumulate as a pool of
free amino acids in the mature endosperm. Accumulation
results in part from lysine degradation being compromized,
because lysine-ketoglutarate reductase activity is suppressed,
but other studies have identified changes in sensitivity of the
monofunctional AK (Ask2) to feedback regulation by lysine as
a major genetic determinant of free amino acid content. In a
mapping population with the mutant Ask2 was a good candi-
date gene for a quantitative trait locus (QTL) affecting free
amino acid content in the harvested grain. Dihydropicolinate
synthase (DHDPS) converts the product of AK (aspartate 4
semialdehyde) to an intermediate in lysine synthesis and is
also subject to feedback inhibition by lysine during seed
development.
Specific features of C metabolism within the developing
seed are required to accommodate some aspects of amino
acid synthesis. Phosphoenolpyruvate carboxykinase (PEPCK)
is present in a wide range of developing seeds. In grape seeds,
PEPCK is expressed in the inner layer of the seedcoat, in the
chalaza and in cells at the boundary of the storage tissue;
expression coinciding with maximum deposition of storage
reserves. Feeding asparagine to developing grape seeds caused
a strong induction of PEPCK indicating that the enzyme func-
tions in the metabolism of N-solutes after unloading from
phloem. These data are consistent with an anaplerotic role
for PEPCK in supplying acetyl-CoA when the TCA cycle is
involved in the interconversion of C skeletons for 4C and 5C
amino acids.
Because seeds are formed within enclosing structures that
limit gas exchange the concentrations of respired CO2 within
the structures may build up to quite high levels and refixation
of some of this C is achieved through RubisCO (Ribulose bis P
carboxylase/oxygenase) and especially PEP carboxylase. In
lupin embryos some of this refixed C is incorporated in sugars
and TCA cycle intermediates (malate, isocitrate, and citrate),
but as much as one-third provides C for a range of amino acids,
principally alanine, aspartate, and serine, but also glutamate,
glutamine, asparagine, glycine, valine, leucine, and isoleucine.
In Vicia faba PEP carboxylase is up-regulated in cotyledons
during grain filling. Again this suggests an anaplerotic role for
the provision of 5C skeletons to the TCA cycle functioning in
amino acid synthesis in an intensely respiring tissue.
Synthesis of Stored N Reserves in Seeds
The major dietary sources of plant protein are seeds of cereals
and legumes. Legumes and some oilseeds typically contain
3050% dry weight as protein while cereals are lower,
10%.
The storage proteins constitute up to 90% of the seeds protein
at maturity and as a consequence it is this group that largely
determines the nutritional and processing qualities of grain.
Features of the synthesis, localization, and properties of the
major storage proteins have been reviewed extensively and,
more recently, details of the regulatory mechanisms that deter-
mine their synthesis in response to changes in plant nutrition
and environmental conditions are being revealed.
While most of the incoming N is used for protein synthesis,
seeds may also accumulate quite a significant pool of free
protein amino acids that persists to maturation and constitutes
a store at maturity. The amides glutamine and asparagine are
common constituents of this soluble-N pool but other amino
acids may predominate in some species.
One of these soluble reserves is arginine. In soybean argi-
nine is synthesized in developing cotyledons as well as being
supplied from the seedcoat to the embryo through the apo-
plast. Its accumulation to more than 60% of the free amino
acid pool apparently occurs because arginase is not expressed

in the embryo. The accumulation of lysine and other amino
acids of the aspartate family in maize endosperm was noted
above and doubtless there are numerous other examples that
occur in nature.
Seeds accumulate significant quantities of non-protein
amino acids. Some of these, such as homoserine, are interme-
diates in the synthesis of the 20 protein amino acids while
others are accumulated as end products of metabolism.
Some, such as the arginine analogue canavanine, are highly
toxic to insects and other potential herbivores apparently serv-
ing to protect the developing or mature grain. Canavanine can
reach levels that are 6% of the dry weight in the seeds of
Canavalia ensiformis and so represent a significant stored
reserve of N. Mature groundnut (Arachis hypogea) seeds contain
high levels of g-methylene-glutamine that is mobilized to the
developing seedling following germination but is not metabo-
lized to any great extent, apparently accumulating again in the
next generation of seed.
There are hundreds of nonprotein amino and imino acids
formed by plants with the overwhelming majority of these
accumulated in seeds. Many of these occur in sufficient
amounts that they could be construed as a stored reserve but
the physiological and biochemical studies to confirm this role
have yet to be undertaken.
Mobilization of N Reserves in Germinating Seeds
While the N reserves in seeds are principally in the form of
storage proteins, as germination proceeds N from nucleic acids
or metabolic proteins that might have been formed before
dehydration or subsequently following imbibition is also
available. The possibility that allantoin is formed from nucleic
acid catabolism has been indicated earlier. Additionally seeds
mature with what can be a significant soluble N pool and this
appears to have a specialized role in the first few hours/days of
germination when the newly activated meristems of the
embryo begin to grow rapidly.
Arginine has been recognized as a common constituent that
fulfils this role. Its accumulation as the seed matures results
from suppression of an enzyme, arginase, which would nor-
mally catabolize the amino acid. However, within a day after
imbibition arginase transcript level and activity increase
sharply in the cotyledons to release urea and, as a result of
urease activity, ammonium ion for reassimilation and translo-
cation to the new meristems. It is likely that the translocated
N-solute at this time is glutamine. Thus arginine is a compact
form of stored N (C:N ratio of 1.5) that provides an almost
instant N source, and is no doubt used in this role in a wide
range of species (e.g., pumpkin and pea seeds). Whether other
soluble N-compounds that accumulate in maturing seeds func-
tion as initial sources of N for germination is not known.
Ammonia released from oxidation of free and mobilized
amino acids can account for 50% of the N made available
during germination of Arabidopsis seeds indicating, as during
seed development, a central role for GS/GOGAT in reassimila-
tion. QTLs for enhanced germination efficiency in maize have
been identified that are co-localized with two of the GS1
(cytosolic forms of glutamine synthetase) structural genes,
gln3 and gln4, emphasizing the importance of ammonia
PROTEINS | Nitrogen Metabolism 127
reassimilation in regulating the germination process. Such
markers offer the potential to breed for complex traits related
to seed N metabolism in grain improvement program.
Future Prospects
Despite the wealth of knowledge about the composition of
seeds and the nature of their stored reserves, factors that deter-
mine which reserves are formed and stored and the roles of
environmental cues in these processes have yet to be estab-
lished. Clearly there is tissue-specific and temporal expression
of genes that encode biosynthetic enzymes but the nature of
the mechanisms that integrate metabolism so that stored
N-reserves are generated to maximize the available resources
of C, N, and S need to be resolved. Although functional geno-
mic or proteomic analyses have been applied to developing
seed tissue they have yet to reveal new information about more
detailed aspects of the pathways and regulation of N metabo-
lism and transport. As genomic analysis is extended to a wider
range of plants this information will become available provid-
ing new tools for both understanding and manipulating grain
development. Significant changes to grain composition
through selection from germplasm collections, conventional
breeding and the exploitation of direct genetic engineering will
be an ongoing area for research as novel, more nutritious and
health promoting food ingredients are sought. Because plants
engage in a bewildering diversity of N metabolic end products,
many of which accumulate in seeds, it is important that critical
studies of N metabolism are extended beyond the handful of
species for which information is currently available.
Exercises for Revision
Draw up a flow chart showing the movement of nitrogen
from the farmers field to the final grain-based food
product.
Investigate further the concepts of source and sink in rela-
tion to carbon and nitrogen metabolism in cereals. How
useful are these concepts?
Exercises for Readers to Explore the Topic Further
Investigate the current understanding of nitrogen metabo-
lism in relation to endosperm protein formation in a
selected cereal grain.
What techniques are available to study nitrogen fluxes in
cereal plants?
What are the energetic limitations to increased protein
content in cereals?
See also: Carbohydrates: Carbohydrate Metabolism; Starch:
Chemistry; Proteins: Protein Synthesis and Deposition; The Enzymes
Associated with Food Grains and Their Functions in Processing; The
Protein Chemistry of Dicotyledonous Grains; The Basics: Grain:
Morphology of Internal Structure; Wheat Processing: The Gluten
Proteins of the Wheat Grain in Relation to Flour Quality.
128 PROTEINS | Nitrogen Metabolism

Further Reading
Editorial (1995) The plant cell special review issue on plant biochemistry. Plant Cell
7: 7951111.
Atkins CA (2000) Biochemical aspects of assimilate transfers along the phloem path:
N-solutes in lupins. Australian Journal of Plant Physiology 27: 531537.
Azevedo RA, Arruda P, Turner WL, and Lea PJ (1997) Review Article Number 121: The
biosynthesis and metabolism of the aspartate derived amino acids in higher plants.
Phytochemistry 46: 395419.
Bewley JD and Black M (1985) Seeds. Physiology of Development and Germination.
NY: Plenum Press.
Kozlowski TT (ed.) (1972) Seed Biology, vol I. Importance Development and
Germination. NY: Academic Press.
Lea PJ and Ireland RJ (1999) Nitrogen metabolism in higher plants. In: Singh BK (ed.)
Plant Amino Acids, pp. 147. NY: Marcel Dekker.
Murray DR (1987) Nutritive role of seedcoats in developing legume seeds. American
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Rosenthal GA (1982) Plant Nonprotein and Imino Acids. NY: Academic Press.
Relevant Websites
mpiz-koeln.mpg.de http://www.mpiz-koeln.mpg.de.
dijon.inra.fr http://www.dijon.inra.fr.
 Nitrogen in Grain Production Systems
XM Zhou, McGill University, Ste-Anne-de-Bellevue, QC, Canada
BL Ma, Agriculture and Agri-Food Canada, Ottawa, ON, Canada
DL Smith, McGill University, Ste-Anne-de-Bellevue, QC, Canada
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 2, pp. 277286, 2004, Elsevier Ltd.
Topic Highlights
The nitrogen cycle refers to the dynamic flow of nitogen
from the atmosphere to plants and animals and its
recycling.
Nitrogen is fixed through nitrogen (atmospheric fixation),
biological fixation, and industrial fixation through the
Haber-Bosch process.
Nitrate and ammonium are taken up and assimilated by
plants, and produce amino acids and thence the storage
proteins of the grain.
Inappropriate use of fertilizers may result in nitrate run-off
to surface water, and consequent environmental health
risks.
Learning Objectives
To understand the fixation of nitrogen from the atmo-
sphere and its processing into storage proteins of the grain
or seed.
To understand the importance of crop management prac-
tices in reducing environmental harm caused by nitrogen
runoff.
Introduction
Nitrogen (N) is the largest component of the Earths atmo-
sphere, comprising
78% (by volume) of the air we breathe.
It is usually present in the atmosphere as a dinitrogen gas (N2),
which is colorless, odorless, tasteless, and relatively inert at
room temperature.
Nitrogen was first identified by the chemist and physician
Daniel Rutherford in 1772. It is now known to be present in a
vast range of materials encountered in our day-to-day exis-
tence; foods, fertilizers, poisons, and explosives are a few gen-
eral examples. Some specific examples are: nitrogen gas is used
as a blanketing medium during the production of electronic
components, as an agent for annealing stainless steel, in bev-
erage processing, as a refrigerant for food protection and pres-
ervation, and as a major fertilizer in crop production.
Atmospheric nitrogen is also responsible for the orange-red,
blue-green, blue-violet, and deep violet colors of the aurora.
More importantly, N is a component of a wide range of
biological compounds, being fourth in abundance after
carbon (C), hydrogen (H), and oxygen (O).
Nitrogen exists in organic and inorganic forms in the atmo-
sphere, biosphere, hydrosphere, and geosphere. It can be pre-
sent in gas, liquid (dissolved in water), or solid phases. The
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00103-0
most common inorganic forms include dinitrogen (N2) gas,
nitric acid (HNO3), nitrate (NO 
3 ), nitrite (NO2 ), ammonium
(NH4 ), gaseous nitrous oxide (N2O), gaseous nitric oxide
(NO), gaseous nitrogen dioxide (NO2), and gaseous ammonia
(NH3). The most common N-containing biomolecules belong
to one of the following three chemical families: proteins
(including enzymes) and related compounds (including pep-
tides and amino acids), nucleotides (largely RNA and DNA),
and porphyins (e.g., cytochromes and chlorophyll).
Although N constitutes only 17% (w/w) of plant dry
matter, it is the element most frequently limiting the plant
growth. Sufficient nitrogen is fundamental for crop growth,
development, and yield. Improvement in N use-efficiency is
thus the key to produce safe and high-quality food and feed in
an environmentally responsible way.
The Nitrogen Cycle
Nitrogen is a dynamic element and is recycled continually by
plants and animals. It can be transformed from one oxidation
state to another through processes that comprise the N cycle
(Figure 1). The term cycle is really a misnomer as the path-
ways of N in the biosphere are actually a simple web. Nitrogen
moves among oxidation states, with ammonia (NH3) being
the most reduced and nitrate (NO 3 ) the most oxidized. The
tremendous chemical stability of dinitrogen (N2) gas makes
the conversion of this compound into ammonia (NH3) or
oxides of N (NO 3 ) energetically expensive and the rate-
limiting step in this part of the cycle. This causes N2 to
accumulate in the atmosphere. Thus, cycling of nitrogen by
various biological entities determines a major part of the com-
position of the Earths atmosphere.
It is a paradox that while 78% of the Earths atmosphere is
N2 gas, and plant stems, leaves, flowers, and fruits are
immersed in this gas, biologically available N is generally the
most limiting nutrient element for crop production. A solution
to this problem evolved in the exquisitely adaptable prokary-
otes. A limited number of microorganisms are able to react N2
with electrons to form NH3, which they then incorporate into
various biomolecules. When these microorganisms die, N is
mineralized to simple forms (NH3, NO 3 ) through decompo-
sition by other microorganisms. In these forms, N is available
to higher plants. Nitrogen is taken up by crop plants, which
become food for animals, including human beings, and energy
sources for bacteria or fungi; N is then cycled back to the soil
when dead cells and tissues of organisms (plant debris for
example) or N-containing waste products (such as animal
urine and feces) are broken down by soil microbes.
129
130 PROTEINS | Nitrogen in Grain Production Systems

Removal N
from harvest

Gaseous losses N2, Chemical

NOx into fertilizer
atmosphere Rainfall (HNO 3) and
fertilizer NH4+ Animal and plant
residues
Denitrification

Soil organic matter

Plant uptake R-NH2
Symbiotic and nonsymbiotic
N2 fixation

Nitrates NO3 Mineralization

Nitrification
Soluble and exchangeable
ammonium NH4+
Micelle fixation
Nitrites NO2 Nitrification

Metal and clay organic

complexes
Leaching

Figure 1 A diagram of the N cycle in a typical grain crop containing agri-ecosystem.

In natural ecosystems, N changes from one form to the
other, but most of it remains within the nearly closed system
in a dynamic balance. In agricultural systems, humans often
add chemical N, fixed through industrial processes, a signifi-
cant portion of which is eventually removed from the field in
harvested material. Food, fiber, and livestock are now often
produced primarily in one location and transported to other
regions of the world for consumption, making crop produc-
tion fields into nutrient flow-through systems. Nitrogen input
into agriculture production systems is largely from chemical
fertilizers, or N-fixing crops (including those grown as green
manures). There are other minor sources, such as biosolid
waste products, which are sometimes added to croplands.
A small amount of N comes with precipitation, sometimes
associated with lightning discharges, which reacts the N2 and
O2 in the atmosphere. There is some N-cycling within agricul-
tural systems, with livestock manures constituting the major
component of this process.
Nitrogen outputs from crop production systems include
removal of harvested grains and forage biomass from produc-
tion systems and losses of various N forms through runoff
(surface water), leaching (to ground water), denitrification
(production of gaseous N2O), and volatilization (generally
NH3). Because of leaching and gaseous losses, and aspects of
plant N physiology, only
50% of the N applied to crop
production systems is taken up by the plants, with the remain-
der being unusable or lost. Globally, various forms of N are in a
dynamic balance.
Soil microbes play a wide range of roles in the N cycle.
Because of this, these processes are dependent on temperature,
moisture, and the quality and quantity of soil organic matter
available to them. It is microbes that break down most of the
organic material, leading to mineralization of organic N.
Microbial populations in the soil can take up large amounts
of N released from decomposing plant tissues, as long as there
is sufficient reduced carbon available, leading to immobiliza-
tion of N. The conversion of ammonium (NH 4 ) to nitrite
(NO 
2 ) and to nitrate (NO3 ) is referred to as nitrification. All
forms of N in the soil (except N2 gas) can be converted to NO 3
by soil microorganisms, under aerobic conditions when soil
temperatures are above freezing.
The amount of nitrification is controlled by the supply of
NH 4 which is, in turn, controlled by decomposition rates,
plant and microbial growth, and soil pH, or by N-fertilizer
addition rates. Nitrate can be converted back into N2, or into
nitrous oxide (N2O), through denitrification. This process
often takes place under anaerobic (generally water saturated)
soil conditions. In this process, soil NO 3 can be reduced to N2
gas through a series of intermediate steps [NO 
3 ! NO2 ! NO
(gaseous nitric oxide) ! N2O] and finally to N2 (gas). The

intermediate and final products of denitrification, are released
into the atmosphere. Denitrification may lead to the loss of
45% of soil NO 3 each day when soil remains water saturated.
Microbes play a key role in most aspects of the N cycle, gener-
ally extracting energy by converting forms of N out of equilib-
rium with the oxidation state of their environment into forms
that are in equilibrium, and extracting a reasonable portion of
the energy liberated through this process. This causes N to
move among forms that are biologically available or unavail-
able, mobile or immobile, oxidized or reduced, gaseous or
solid.
Nitrogen Sources
Nitrogen exists in inorganic or organic forms in our environ-
ment. Inorganic forms include NO 
3 , NO2 , NH3, and N2
whereas organic N is found in a wide range of biomolecules.
Protein is one of major classes of biomolecules and generally
the largest repository of organic N in the biosphere. RuBisCo,
the plant protein responsible for the uptake of CO2 by C3
plants, is the most abundant protein on the planet. The other
two most abundant classes of biocompounds, carbohydrates,
and lipids, generally have little N associated with them. Over
9099% of N in the surface (or tillage layer; 025 cm) layer of
most soils is in organic forms. Although there is an abundant
potential supply of N in the Earths atmosphere (N2 gas), the
unique geometry of the triple bond between the two N atoms
results in a very stable compound that is very energetically
expensive to break. Thus, this form of N cannot be used directly
by most organisms. Whenever dinitrogen is converted into
other forms, a large input of energy is involved. The major
sources of this energy are: (1) lightning in the case of atmo-
spheric fixation, which accounts for 58% of total N fixed and
potentially available to the biosphere. This process results in
the combination of oxygen from the atmosphere with dinitro-
gen, causing the formation of N oxides. These compounds
dissolve in rain water and are carried to Earth; (2) reduced
carbon and adenosine triphosphate (ATP) in the case of bio-
logical N fixation, conducted by certain prokaryotes alone or in
symbiotic relationships with plants and sometimes animals;
and (3) fossil fuels for industrial fixation.
Industrial fixation relies on the HaberBosh process, devel-
oped during the First World War for the production of explo-
sives. This process uses a catalyst, high pressure, and high
temperature (600  C), to combine N from the atmosphere
with hydrogen (usually derived from natural gas or petroleum)
to produce ammonia (NH3). Ammonia can be used directly as
fertilizer, reacted with CO2 to form urea ((NH2)2CO), and/or
oxidized to form nitrate (NO 3 ) which can be salted with other
compounds (such as ammonium, NH 4 ) to form other types of
N fertilizers (e.g., ammonium nitrate). The large requirement
for energy makes N fertilizer easily the largest component of
fossil fuel consumption in crop production, being close to 50%
of the total for crops such as maize.
The bulk of the soil N in noncropland systems comes
from fixation by symbiotic bacteria (largely genera associated
with legumes and collectively referred to as rhizobia and spe-
cies of Frankia, usually symbiotic with woody plants), fixation
by free-living bacteria and mineralization of organic matter
PROTEINS | Nitrogen in Grain Production Systems 131
(N released as decomposition of plant and animal residues).
The potentially available N (9099%) in the soil is in organic
forms, the rest being in inorganic (or mineral) forms. The
organic forms are not immediately available for plant use and
have to be decomposed by soil microorganisms through min-
eralization. In addition, a very small amount of organic N may
exist in relatively small soluble compounds, such as urea or
amino-sugars, which may be slightly available to plants.
Symbiotic (largely legume) N-fixation and mineral fertilizer
N are the major sources of N for grain production systems, with
animal and green manures, crop residues, composts, sludges,
soil micro- and macro-biota being the other possible sources.
Again, organic N must be mineralized before it is available to
plants. Sodium nitrate (NaNO3) and potassium nitrate
(KNO3) are formed by the decomposition of organic matter.
In certain dry areas of the world, these saltpeters are found in
quantity and are used as fertilizers. Other inorganic N com-
pounds are ammonia (NH3), nitric acid (HNO3), the N oxides
(NO, NO2, NO3, N2O), cyanides (CN), etc. However, inor-
ganic N in soil generally comprises only a small portion of
soil N. Hence, despite N being one of the most abundant
elements on the biosphere, N deficiency is probably the most
common nutritional problem affecting crop plant production
worldwide.
Nitrogen Availability to Crop Plants
The availability of N sources is a fundamental requirement for
metabolism as it allows N assimilation into amino acids (for
which photosynthetically produced carbon compounds are
also required) and their availability for protein synthesis. An
adequate supply of N allows leaf growth and photosynthesis.
Inorganic (or mineral) forms of N, such as NH 
4 and NO3 ,
comprise the majority of plant-available N. Other forms of N
must be converted to one of these compounds by either natural
or artificial means before they can be utilized by plants. When
N is first released from organic matter, it is in the form of
gaseous NH3, which is volatile and can be lost to the atmo-
sphere. When NH3 dissolves in water, it acquires a proton and
becomes ammonium (NH 4 ), which carries a positive charge
and is attracted by the soil clay colloids and soil organic matter,
which carry negative charges. Once attached to the soil matrix,
NH 4 becomes part of the cation-exchange process whereby
plants exchange a hydrogen ion (H) for one of the positively
charged molecules in the soil. Only a small portion of the soil
NH
4 exists in the soil solution; this NH4 is available to crop

plants. On the other hand, NO3 carries a negative charge, and
so is not bound by clay particles. Thus, NO 3 molecules are free
to move with the soil water and are more readily available to
crop plants than NH 4 . However, in coarse textured or water-
saturated soils, NO 3 is much more prone to leaching into
groundwater than NH 
4 ; once in ground water NO3 becomes
a potentially serious pollutant.
Nitrogen Fixation
A symbiotic relationship between leguminous plants and
N-fixing bacteria allows for plant-based biological N fixation.
132 PROTEINS | Nitrogen in Grain Production Systems
The major conversion of N2 into NH3 (a biologically available
form) is the biological N-fixation process, carried out by single-
celled prokaryotes. This process requires a great deal of energy,
supplied as ATP, which is derived from the breakdown of
organic matter, or through photosynthesis (e.g., cyanobacteria).
It is estimated that biological N fixation accounts for the annual
production of as much as 250  109 kg of ammonia, which is
twice the amount produced by the HaberBosch process.
Biological N fixation can be described as the following
equation:
N2 8H 8e 16ATP 2NH3 H2 Leaching
16ADP 16Pi
This reaction is conducted by prokaryotes only (eubacteria
and archea) through the enzyme dinitrogenase, and the closely
associated dinitrogenase reductase. Dinitrogenase contains
both iron and molybdenum in a cofactor referred to as FeMoco.
The fixation of N is carried out while N2 is bound to dinitro-
genase. Dinitrogenase reductase (the Fe protein) is reduced by
electrons donated by a protein generally containing ferredoxin.
Reduced dinitrogenase reductase (Fe protein) binds ATP and
reduces dinitrogenase (molybdenum-iron protein), which pro-
vides electrons to N2 resulting in its reduction to 2NH3.
Due to its economic importance and agronomic value, the
legume-rhizobia symbiosis is the most investigated process of
biological N fixation. Many grain legumes are important crops
due to this symbiotic relationship; these include soybean,
bean, mung bean, faba bean, peanut, and pea as well as forage
legumes such as alfalfa, clover, sainfoin, fenugreek, and vetch.
During the establishment of this symbiotic process, rhizobia
infect the roots of legume plants and cause the formation of
nodules. It is in these nodules that the bacteria fix N and supply
it to the plant. The plant provides reduced carbon (sugars) to
the nodules, where it is converted to organic acids and supplied
to the bacteria as an energy source. In most cases, the symbiosis
can supply all N required for normal growth and development
of the plant. The amount of N fixed by legume plants is
estimated to range from 11 to 250 kg N ha1 year1. When
fertilizer N is supplied, N fixation is inhibited through several
long and short-term mechanisms, so that adding N fertilizer to
N-fixing legumes does not increase the amount of N available
to the plants. For grain legumes, however, a small amount of
starter N fertilizer greatly helps crop growth prior to the time
when nodules become functional and begin to supply N. The
simplest approach for evaluating whether nodules in field-
grown legumes are able to fix N is to observe their interior
color: effective nodules contain leghemoglobin and are bright
red, while nodules not yet functional are generally white and
those that are no longer functioning are generally greenish.
Leghemoglobin carries oxygen into N-fixing (rhizobia contain-
ing) nodule cells, ensuring the availability of large quantities of
oxygen, but in a very controlled fashion, to the nodule-
enclosed rhizobia.
On the other hand, all field crops in the grass family, such as
the grain crops maize, sorghum, rice, and wheat, as well as forage
grasses, and also nonleguminous broadleaf field crops (e.g.,
sunflowers, potatoes, sugar beets, cotton, etc.) are unable to
form N-fixing root nodules. Therefore, these plants must obtain
their required N from the soil and other exogenous sources.
Table 1 Annual N inputs and outputs of typical agricultural land
N (kg ha1)
Inputs
Plant residues 10100
Biological N fixation 10250
Natural fertilization through precipitation 050
Chemical fertilizer 0400
Outputs
Volatilization of NH3 060
Dinitrification 1070
1070
Crop harvest 10150
At present, almost 90 genera of diazotrophic bacteria,
which can reduce atmospheric dinitrogen to ammonia with
varying efficiencies, are known. Biological N fixation can be
carried out by a wide range of prokaryotes, alone or in symbi-
osis, including cyanobacteria free living and in symbiosis in
lichens, cycades, the fern Azolla or the angiosperm Gunnera;
and free-living soil bacteria. These types of N fixation contrib-
ute significant quantities of NH3 to natural ecosystems, but not
to most cropping systems, with the exception of paddy rice.
Their contributions are generally less than 5 kg N ha1 year1.
Table 1 indicates the general annual input and output of N in
agricultural ecosystems.
Symbiotic Signals
Nitrogen fixing root nodules are the result of symbiotic inter-
actions between leguminous plants and rhizobia. This begins
with a recently elucidated signal exchange between the two
symbiotic partners. Initially, the roots of leguminous plants
secrete flavonoid compounds into the soil. These compounds
activate a set of genes in the appropriate rhizobia. A struct-
urally diverse mixture of lipo-chitooligosaccharides (LCOs) is
produced by rhizobia after induction with flavonoids. LCOs
consist of a chitooligosaccharide backbone of b-1,4-linked
N-acetyl-d-gluconsamine (GlcNac) and a fatty acyl group
attached to the nonreducing saccharide. Diversity of the fatty
acyl substituents (length and degree of unsaturation) contributes
to the diversity of LCOs. In addition, various groups can be
added to the chitin backbone. The nodE gene determines
the nature of the fatty acyl moiety, a major determinant of
host range. LCOs are considered to be key factors in the specific
recognition processes that underlie the formation of legumi-
nous root nodules and their bacterial infection; in these asso-
ciations only appropriate legume and rhizobium matches can
come together and create a specific symbiosis.
Uptake
Nitrate (NO
3 ) and ammonium (NH4 ) are the two major
mineral forms of N taken up and assimilated by plants.
These compounds enter the plant roots by diffusion through
the space between root cells until they cross the plasma lemma
and enter the plant cell interior. Once inside the symplast

(for a review of the symplast, see www.worldreference.com)
these compounds can be stored in vacuoles, assimilated into
amino acids or proteins or moved into the stele (xylem) for
transport elsewhere in the plant. Movement among the cells
that make up the symplast is through the plasmodesmata that
connect plant cells. All mineral N must enter the symplast
before entering the stele. This can happen at any point in the
root cortex, but must happen by the time mineral N molecules
reach the casparian strip. During the course of the vegetative
growth, both NO
3 and NH4 enter the plant root and are
transported via the xylem to the leaf (Figure 2). Once it
reaches the leaf, nitrate reduction takes place, leading to the
production of amino acids. Most of the newly formed organic
acids are then translocated to the root where carboxyl groups
are exchanged for an ammonium and the newly assimilated
N is incorporated largely into leaf N compounds (protein).
When a large quantity of N is required by developing fruits
and filling seeds (reproductive growth stage), leaf proteolysis
occurs and a great deal of the amino N in the leaf is ultimately
exported to the filling fruits where it is accumulated as
seed storage proteins (Figure 2). As a result, the phloem is
enriched with amino compounds, which repress nitrate uptake
and diminish the rate of nitrate reduction. Improved crop
yield due to increased fertilizer N application suggested that
the availability of N (usually nitrate in agricultural soils) is
often the limiting factor in plant growth. The absorption of
NO 3 by roots is not only determined by the N demand of
the plant, but also by its availability in the soil environment.
Nitrate absorbed via roots is either reduced in situ into
ammonium, imported into vacuoles (by high- and low-affinity
transport systems) for storage, or transported via the xylem to
the shoot where it can either be metabolized or stored as a
reserve. A low-affinity root NO 3 uptake system may play a
greater role in nitrate uptake. In some cases, a high-affinity
system has been shown to be the major mode of nitrate uptake
through uptake competition studies and membrane voltage
change studies. The low affinity transport system (LATS)
(possibly a carrier system or an anion channel), and the
inducible, high affinity transport system (HATS) are regulated
by cellular energy supply, intracellular nitrate levels, and a
proton electrochemical gradient. The latter is regarded as an
H/anion cotransport carrier system that produces transient
plasma membrane depolarization upon addition of nitrate.
The depolarization is counteracted by the plasma membrane
H-ATPase. The plasma membrane proton ATPase is induced
by nitrate. In addition to the nitrate uptake system, plants
have an inducible nitrate efflux system, requiring both RNA
and protein synthesis. The efflux system, however, has a much
slower rate than the uptake system.
In the agricultural soils of tropical and temperate regions
(except paddy rice fields), nitrate accounts for 7090% of soil
mineral N. Nitrate in the soil solution is 10 times more mobile
than NH
4 . Two reserves of soil NH4 exist: a small portion of
NH 4 is in the soil solution, and a large portion of NH 4 is
adsorbed on soil colloids. Only water soluble NH 4 is available
for plant uptake. Depending on the plant species, the bulk of
the nitrate entering the plant is reduced to ammonia and
assimilated into organic compounds in the roots or in the
leaves. Plants can take up either NO
3 , NH4 , or mixtures of
the two. In most cases, crop plants perform better when the
PROTEINS | Nitrogen in Grain Production Systems 133
mixture of NO
3 and NH4 fertilizer is provided than either one

alone. Uptake of NO3 by roots is governed by the concentra-
tion of NO 3 in the soil solution, the volume of soil exploited
by roots and root density within that volume, and plant
demand for N, the latter being largely related to plant growth,
development, and production. The root uptake efficiency of N
is affected by metabolic demand, N concentration around the
roots, and conditions such as temperature.
Ammonium is mainly assimilated in roots. It or alanine,
formed by organic acid ammonization, is the main form of N
exported from symbiotic N2-fixing microorganisms to their
host plants. In soil, NH 4 comes mainly from the mineraliza-
tion of organic matter. When NO 3 levels in the soil are low
or completely exhausted, plants can rely on NH 4 released
from cation-exchange sites or through mineralization of
organic matter. During NH 4 assimilation in the root, excess
H is generated (3NH
4 3R  NH2 4H ) and released

directly to external solution. While NO3 assimilation occurs
excess OH is produced (3NO 
3 3R  NH2 2OH ). This
excess OH may be neutralized by carboxylation or by
HCO3 released from the roots. Thus an H neutral synthesis
of plant materials can occur with a ratio of assimilation of
1NH 
4 per 2NO3 . This is why many plant species grow opti-
mally with a mixture of mineral N compounds (NO 3 and
NH 
4 ). With a mixed N solution (NO3 and NH4 ), NH4 is
often the form of N preferentially taken up by the plant.
Many forest species take up primarily NH 4 , since forest soils
are relatively acidic, inhibiting the development of popula-
tions of bacteria that cause nitrification. The average NH 4
concentration in well-aerated soil is 10100 times lower
than NO 3 , rarely exceeding 0.12 ppm. However, absorption
of NH 4 by plant roots can occur at very high rates, due to the
presence of transport systems in the root plasma membrane
with high substrate affinities. Like NO
3 , NH4 uptake by roots
is diurnally regulated, with maximum uptake during the light
period. Uptake is stimulated by sugar supply, implying that
the greater uptake during the day is related to the availability
of photosynthate.
Assimilation of Nitrogen
Both nitrate and ammonium are taken up and assimilated by
plants. In general, NO 3 is the predominant form of inorganic
N taken up by grain crop plant roots (except paddy rice).
Under most circumstances, absorbed nitrate is reduced to
ammonium in roots and/or leaves, and is then utilized in the
synthesis of amino acids and proteins. Whatever the source of
ammonium in plants, this ion is first converted into amino
acids, the building blocks of protein.
Nitrate is assimilated into organic compounds by GS/
GOGAT (GS glutamine synthetase/glutamate synthase;
GOGAT glutamate oxidase, glutamine amino tranferase).
The GS/GOGAT pathway is considered the major pathway for
incorporating reduced N into organic molecules. This pathway
recycles glutamate; amino groups are eventually transferred to
other amino acids and utilized for protein synthesis and also
for synthesis of other key biomolecules such as RNA, DNA, and
porphyrins.
134 PROTEINS | Nitrogen in Grain Production Systems

Ammonia assimilation

Proteins Proteins
Seeds
Mitochondrion Chloroplast
AA
Albumins NH1
Globulins GS2 NIR
NH4+ NO2
Prolamins NH1
(cereal) Vacuole
NH4+
NH4+

NH+4 HNO3
Leaf
Stem
NO3 NO2

Phloem
Nitrate reduction
(cytoplasm)
NR
HNO3
NO3 NO2
NH+4

NO3 Nitrite reduction

(plastid)
NO2 NIR NH+
4
Xylem
Amino acid
Protein
(AA)

Root Symbiotic nitrogen fixation

NH+4 NO3
N2 + 3H2 2NH3
HNO3

Plasma membrane
Nitrate reduction
AA GS2 Plastid (cytosol)
AA NIR
+
NH4 NO2
GS1
Vacuole NR
NH+4 NH+4 NH+4 NO3 HNO3

NH+4 NO3

Figure 2 A diagrammatic summary of N metabolism in a grain crop.

GS and GOGAT are located in the cytosol (for a review of
the cytosol, see www.worldreference.com) and the chloro-
plast (for a review of chloroplast, see www.worldreference.
com). Their activity in particular plant tissues is closely linked
to specific primary N assimilation, NH3 recycling during pho-
torespiration, or general N remobilization. There are two path-
ways for incorporation of N into amino acids: (1) Direct
incorporation where glutamate dehydrogenase is involved:
a-ketoglutarate NH3 NADH ! glutamate NAD; (2) GS-
GOGAT cycle: GS catalyses the ATP-dependent conversion of
glutamine utilizing ammonia as substrate and is represented by
two protein groups, plastid (GS1) and cytosolic (GS2) isoen-
zymes; GS1 is located exclusively in chlorophyllous tissues while
GS2 is located predominantly in roots, nodules, and floral

organs. GOGAT catalyzes the conversion of glutamine and
a-ketoglutarate to glutamate: glutamine a-ketoglutarate
NADP H H ! 2 glutamate NADP. The assimilation of NH 4
NH3 may be by either glutamate dehydrogenase or GS/GOGAT;
the first of these requires quite high concentrations and the
bulk of uptake is thought to be through GS/GOGAT.
Following the incorporation of NH3 into glutamine, N can
be transferred directly to the same position in asparagine by
asparagine synthetase. Amino transferase can transfer the
amino group to aspartate, which is converted to asparagine. It
has been suggested that the reaction catalyzed by Fd-GOGAT is
the key regulatory element controlling N partitioning and
redistribution during plant growth and development. To
date, research has focussed on modifying ammonia assimila-
tion in transgenic plants and this has demonstrated that the
amplification or shifting of ammonium assimilation in a par-
ticular organ or tissue has strong effects on plant growth and
development. Thus, there appears to be great potential for
improving N-use efficiency in crop plants.
Reduction of Nitrate and Nitrite
Once NO 3 enters the root symplast, it may follow five
pathways:
1. efflux back to the apoplast [for a review of apoplast see
www.worldreference.com];
2. reduction of NO 
3 to NO2 , then NH4 by nitrate reductase
(NR) using the reduced form of nicotinamide adenine
dinucleotide (NADH) and by nitrite reductase (NIR) using
ferredoxin (Fd) in root cells, leading to the production of
amino acids;
3. accumulation into vacuoles of root cells, involving trans-
port across the tonoplast [for a review of tonoplast see
www.worldreference.com];
4. secretion into xylem vessels for long distance transport to
the shoots; and
5. reduction of NO 3 in the leaves, often using excess reductant
in upper, CO2 limited, leaves.
High concentrations of nitrate can be found in vacuoles, indi-
cating that nitrate not only acts as a nutrient but also partici-
pates in osmotic maintenance. The first step in the nitrate
assimilation pathway is the reduction of nitrate to nitrite,
catalyzed by assimilatory NR. When reduction occurs in leaves,
the nitrite formed by NR activity is then translocated to the
chloroplast, where it is further reduced to NH 4 by NIR. During
the nitrate reduction process, NR activity appears to be the rate-
limiting step in the conversion of NO
3 to NH3/NH4 , because

it is (1) the first enzyme in the pathway NO3 of assimilation,
(2) substrate inducible, (3) relatively unstable, and (4) its
activity relative to other enzymes in the NO 3 assimilation
pathway is low and its Km (related to binding affinity) for
NO 3 is high. Further, there is evidence that when nitrate is the
dominant N source, nitrate reductase activity (NRA) is the
limiting factor to the growth of many plants. NR has therefore
been intensively studied in order to understand its catalytic
efficiency and regulation. Nitrite is reduced to NH 4 by NIR
PROTEINS | Nitrogen in Grain Production Systems 135
using ferredoxin (Fd). Nitrite reductase is located in plastids
and possesses FeS and heme cofactors. Subsequently, the
is incorporated into carbon compounds: amides
(amino acids) and ureides.
Nitrate reduction occurs mainly within the chloroplasts of
green leaves, but can also occur within plastids of roots. Induc-
tion and the rate of nitrate uptake depend on the external
nitrate concentration, light, pH, temperature, the concentra-
tion of other ions, anaerobic conditions, metabolic inhibitors,
and biological genotypes. Since nitrate reductase contains Mo,
Mo deficiency can cause N deficiency.
Nitrogen Storage
Seed storage proteins comprise 70% (up to 100% in some
developing countries) of total intake of dietary protein by
humans. It also provides the major protein source in the diet
for nonruminant livestock. The amino acids in the leaves,
produced from assimilated NO 3 with energy and carbohy-
drates produced by photosynthesis, are transported to the
embryo and the cotyledons via mass flux in phloem vessels
of the leaf. The main transport forms of amino acids are:
glutamine, asparagine, serine, alanine, glutamate, and aspar-
tate. As a result, storage proteins are the products of the
secretary pathway, which resides within the endomembrane
system of the cell. In general, N assimilates available to
developing grains are mainly used for the synthesis of pro-
teins. These proteins then are accumulated and stored in
specialized tissues such as the endosperm of cereals, other
endospermic monocotyledons, or the cotyledons of the
embryo for the pulse legumes and other nonendospermic
dicotyledons.
Grains of cereal crops usually contain 719% protein while
the seeds of legume crops contain up to 40% protein, in both
cases on a dry matter basis. Tuber crops are rich in soluble
carbohydrates and low in protein. For example, the crude
protein concentration in sweet potato is twice that of potato
(12.1% versus 5.4%). Thus, when tuber crops are used as
human food or animal feed, they are a good source of energy,
but require protein supplementation.
Almost all storage proteins in the seeds of major crops can
be grouped into three categories according to their functions:
(1) globulins (soluble in dilute saline solutions), the most
common storage proteins, are present in all angiosperm
seeds; (2) albumins (soluble in water); (3) prolamins (soluble
in dilute alcohol) and restricted to the seeds of Gramineae
(grass) family. This protein represents the main storage protein
in cereals such as wheat (gliadins), barley (hordeins), oat
(globulins), rice (glutelins), rye (secalins), and maize (zeins).
Both albumins and globulins have biological functions. This is
particularly true of enzymes during seed germination. In most
cases, storage proteins have no biological activity and only act
as a source of N, sulfur, and carbon skeletons for the develop-
ing seed. Thus, storage proteins are not as evolutionarily con-
strained as those of other proteins, such as enzymes, although
there is a requirement that proteins should be efficiently syn-
thesized, packaged, and stored, and then remobilized during
the process of seed germination.
136 PROTEINS | Nitrogen in Grain Production Systems
Interactions between Nitrogen and Carbon Metabolism
The products of photosynthesis are the building blocks of
agricultural production. From a production point of view, the
efficiency of photosynthesis is mainly determined by the avail-
ability of light, CO2, water, heat (temperature), and key ele-
ments in the soil, with N being the most likely to be limiting.
Precipitation is outside management control in crop produc-
tion systems, although irrigation may provide reliable water
where water resources are available. Nitrogen application,
including aspects such as quantity, timing, and method is
part of most crop management regimes. Grain yield is largely
a function of accumulation of photosynthetic assimilates. The
interactions between carbon dioxide (CO2) and nitrate assim-
ilation and their dynamics are key elements of crop produc-
tion. As a result, an adequate supply of N, its assimilation into
amino acids (for which photosynthetically produced carbon
compounds are required), and their availability for protein
synthesis, are essential for metabolism.
Nitrate is reduced to NH3 by NR and NIR using electrons
from photosynthetic electron transport. The NH3 is then con-
verted into amino acids by the GS/GOGAT reaction (the main
pathway by which plants convert ammonia into amino acids),
and the carbon-skeletons are provided as organic acids
derived from the tricarboxylic acid cycle. Carbohydrates for
organic acid synthesis, ultimately from photosynthetic CO2
assimilation, and ATP for the GS/GOGAT reaction are pro-
duced by photosynthesis and respiration. Thus, there is close
interaction in the very earliest phases of N and carbon metab-
olism, both using light energy, with some 10% of the electron
flux in photosynthesizing leaves used for N reduction.
During the last 46 decades grain yields of major field
crops have increased
2.5 fold, largely due to an increase in
the production and storage of carbon in grains by efficient
crop varieties and increased use of N fertilizer. Although large
variations in grain protein concentrations exist among varie-
ties within a species, an increase in yield has generally
resulted in a slight to severe decrease in the protein/starch
or oil ratio. For example, in the case of maize, the increase in
the genetic-based yield potential after 1967 was, on average,
accompanied by a decrease in the concentration of protein
(
1.5%) and an increase in that of starch (
2%). In the case
of soybean, genetic selection leading to a seed yield increase
of 100 kg ha1, increased seed oil yield by 1 kg ha1, and
decreased protein yield by 23 kg ha1. For wheat, an
increase in yield of
2 kg ha1 led to a decrease in the
protein concentration of more than 2% of the dry weight.
Plant geneticists and breeders continually make efforts to
break this relationship, hoping for increases in yield without
decreases in quality.
Nitrogen and Environmental Health
Nitrate accumulation in plants occurs when the speed of
nitrate uptake exceeds the rate of reduction and subsequent
assimilation into amino acid and proteins for growth. A high
nitrate accumulation in edible parts of plants consumed as
vegetables is a potential health hazard. Although nitrate itself
is not toxic, it can easily be reduced, internally, to the toxic
compound nitrite. Reduction to nitrite in vegetables can occur
postharvest as well as after ingestion, in saliva and in the
gastrointestinal tract. One of the symptoms of nitrite toxicity
is methemoglobinaemia, in which NO 2 binds tightly to hemo-
globin reducing the ability of the blood to carry oxygen and
leading to respiratory dysfunction. Chronic nitrite toxicity may
lead to the formation of carcinogenic nitrosamines. The fre-
quency of gastric cancer could be reduced by avoiding high
intake of nitrate.
According to the International Fertilizer Association,
annual world consumption of N fertilizer is equivalent
to
83 million tonnes (Mt). North America accounts for 15%
of this total world consumption. Nitrogen that is not taken up
by plants may lead to gaseous loss to the atmosphere, or an
increased level of nitrate in the soil. This nitrate is vulnerable to
run off in surface water or leaching into groundwater, causing
significant environmental pollution. High concentrations of
NO 3 in drinking water due to surface and ground water con-
tamination can also lead to methemoglobinaemia.
Nitrous oxide (N2O) is a greenhouse gas with a heat-
trapping capacity that is
310 times greater than that of CO2.
Since the last century N2O emission has been increasing 0.3%
year1. Emissions of N2O from cultivated lands are estimated
at 3.03.5 Mt N2O-N year1. The bulk of current N2O emis-
sions are caused mainly by the application of fertilizer N and
biomass burning. When soils become anaerobic, or sometimes
during the conversion of NH 
4 to NO3 , N2O is produced.
N2O escapes the soil as a gas and accumulates in the atmo-
sphere where it is both an agent of stratospheric ozone
destruction and a very potent greenhouse gas. High levels of
soil organic matter and nitrate combined with low oxygen
levels promote rapid denitrification. Increasing emissions
of greenhouse gases are likely to accelerate global climate
change. Average global surface temperature is expected to
rise 0.62.5  C (33.136.5  F) in the next fifty years, and
1.45.8  C (34.542.4  F) in the next century.
Reducing the contamination of drinking water associated
with excess application of inorganic fertilizer and manure in
agriculture production is a high priority. There is an urgent
need to develop new crop production systems that maximize N
use efficiency and are less hazardous to the environment.
However, the reduced application of fertilizers conflicts with
the need for greater crop yields to feed rising world popula-
tions. Crop yields would drop to a very low level if N-fertilizer
applications were to cease. Many countries that use substantial
amounts of fertilizers and export agricultural produce would
barely be able to feed their own population in the absence of
N-fertilizer applications. To overcome the potentially danger-
ous social problems of food shortage, environmental degrada-
tion and pollution, agricultural yields per unit land area must
be increased at the same time as dependence on applied fertil-
izers is decreased. Agriculture uses N fertilizer very inefficiently
(even in developed countries), in part because other environ-
mental conditions are often limiting. Thus, understanding the
process of N uptake and assimilation, and mechanisms associ-
ated with increased N-use efficiency by crops would be a ben-
efit not only in the context of environmental concerns but also
crop quality. Consequently, selection of superior genotypes
with efficient N-use and development of sustainable
 PROTEINS | Nitrogen in Grain Production Systems 137

production systems for specific regions will solve or at least Exercises for Revision
reduce the negative impact of fertilizer N use on the
environment. Compare nitrogen use in leguminous and in cereal grain
crops.
Explain the environmental effects of large-scale nitrogen
Conclusions and Prospects use in crop agriculture.

Nitrogen is essential to the growth and reproduction of all

plants, and animals. Improved understanding of N dynamics
in the biosphere is of both economic and environmental Exercises for Readers to Explore the Topic Further
importance in developing sustainable N management strate-
gies for crop production. Absorption of nitrate and ammo- What is the current supply situation for nitrogenous fertil-
nium by plant roots is the primary pathway for the entry of N izers? What are the constraints to increasing production
into our food chain. Nitrate and ammonium enter plants capacity?
through the cell walls and root membranes. This mineral N How can nitrogen be used more efficiently in agriculture?
has to be incorporated into carbon-nitrogen compounds in How can waste be reduced? How important is precision
order to build cellular metabolites, especially protein. Some agriculture to the development of solutions in this regard?
leguminous plants can convert atmospheric N to bio-available
forms through a symbiotic relationship with specific soil bac-
See also: Agronomy of Grain Growing: Organic Grain Production
teria. This rhizobia-legume symbiosis plays a major ecological
and Food Processing; Carbohydrates: Starch: Synthesis;
and economical role on a global scale. A major recent achieve-
Processing of Grains: Cereals: Breakfast Cereals; Proteins:
ment in understanding the rhizobia-legume symbiosis has
Nitrogen Metabolism; Protein Synthesis and Deposition; The Protein
been the characterization of signal molecules produced during
Chemistry of Dicotyledonous Grains.
initial communication between leguminous plants and
N-fixing bacteria. The specific lipo-oligosaccharide (LCO)
signal compounds, also called Nod factors, secreted by rhizo-
bia, are considered to be a new class of growth regulator, Further Reading
affecting plant growth and development and ensuring the
formation of N-fixing nodules. Consequently, a better under- Follett RF and Hatfield JL (eds.) (2001) Nitrogen in the Environment: Sources,
standing of the whole plant system, from genes, to plant- Problems, and Management. New York: Elsevier.
Lea PJ and Morot-Gaudry JF (eds.) (2001) Plant Nitrogen. Berlin: Springer.
microbe relationships, to biomass production, partitioning
Morot-Gaudry JF (ed.) (2001) Nitrogen Assimilation by Plants: Physiological,
and grain yield formation might achieve the long-term objec- Biochemical and Molecular Aspects. Enfield, NH: Science Publishers.
tive of improved crop N-use efficiency, leading to increased
grain crop yield with reduced impact on the environment. For
cereals and other nonleguminous grain crops, development of
superior genotypes and sustainable production systems with Relevant Websites
efficient N use continues to be one of the major goals for sws.uiuc.edu http://www.sws.uiuc.edu Illinois State Water Survey.
plant geneticists, crop physiologists, agronomists, grain crop fertilizer.org http://www.fertilizer.org International Fertilizer Industrial Association.
producers, environmental scientists, and policy makers. hort.purdue.edu http://www.hort.purdue.edu University of Purdue.
 The Enzymes Associated with Food Grains and Their Functions in Processing
DJ Mares and K Mrva, University of Adelaide, Glen Osmond, SA, Australia
GB Fincher, Australian Research Council Centre of Excellence in Plant Cell Walls, University of Adelaide, Glen Osmond, SA, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Enzymes involved in the synthesis and degradation of
endosperm cell wall polysaccharides.
Synthesis of enzymes during grain development and germi-
nation that are involved in the breakdown of starch and
protein stored in grain endosperm tissue.
Enzymes present in ripe grain that affect the quality,
storage, or nutritional value of wheat and wheat-based
end products.
Endogenous enzyme inhibitors.
Learning Objective
To achieve a better understanding of the role of specific
enzymes in the development, utilization, and storage of
cereal grains.
Introduction
Enzymes are biological catalysts that direct complex biochem-
ical reactions at temperatures relevant to living organisms and
the environments in which they exist. The proteinaceous
nature of these molecules allows for a myriad of three-
dimensional (3-D) structures that will accommodate different
substrate specificities and respond to the presence of other
regulatory molecules, changes in ionic environment, pH, tem-
perature, and hydrophobicity. These interactions often result
in alteration to the conformation of the enzyme protein that in
turn impacts on binding of substrates, activators, inhibitors,
and cofactors or the efficiency of the catalytic activity. In addi-
tion to the mechanisms for controlling enzyme activity, the
actual amount of enzyme protein in a cell can be controlled via
a balance of synthesis and degradation, thus allowing for many
tiers of regulatory control.
Cells in grains contain a multitude of enzymes that are
common to all living cells, including animals and microorgan-
isms, albeit with minor variations in structure and specificity
depending on the species. These enzymes, which are essential
for general cellular metabolism and function, will not be dis-
cussed here, and the reader is referred to any of the numerous
plant biochemistry and general biochemistry texts that are
readily available. Rather, this article focuses on some of the
enzymes that are critical to the specific functions and structures
of cereal grains or those that influence the end use of cereal
grains or the quality of cereal end products. One of the major
functions of cereal grains is to synthesize and sequester starch,
protein, and lipid in the endosperm tissue, which can subse-
quently be mobilized in the germinated grain to support
growth of the developing seedling. In this article, discussion
138 Encyclopedia of Food Grains, Second Edition
of enzyme activities in cereal grains will be presented in the
context of synthesis and degradation of endosperm cell walls in
relation to the specific functions of this specialized storage
tissue and synthesis by grains of enzymes that affect processing,
end-product appearance, nutritional value, or storage of grain.
Enzymes Involved in the Synthesis and Degradation
of Endosperm and Aleurone Cell Walls
Cell Walls in Cereal Endosperm
Cell walls are central to the growth and development of the
endosperm of cereal grains and the remobilization of the
stored reserves during germination. They are also important
contributors to many quality characteristics of cereal grains.
The enzymes responsible for the synthesis and degradation of
cell wall polysaccharides are therefore critical not only to the
normal biological functions of the endosperm but also to the
subsequent utilization of cereals by humans. The endosperm
consists of two distinct tissues, the aleurone and the starchy
endosperm, that have different roles. The starchy endosperm
forms the bulk of the grain. It is the storage organ for starch,
protein, and some lipids, and by the time grains are ripe and
ready for harvest, it is a nonliving tissue incapable of synthe-
sizing new enzymes. By contrast, the aleurone is the layer of
cells that surrounds the starchy endosperm and responds to
signals from the germinating embryo to synthesize and secrete
the key hydrolytic enzymes required to initiate degradation of
the stored endosperm reserves. The aleurone remains a living
tissue throughout development and long periods of storage
and also retains some integrity during the early stages of ger-
mination to enable the programmed dissolution of the endo-
sperm to proceed. Subtle variations in the fine structures of
major wall polysaccharides can have a dramatic effect on the
functional and rheological properties of cereal-based products.
The major wall components in the endosperm of cereal grains
are arabinoxylans and (1,3;1,4)-b-glucans; other wall compo-
nents include cellulose, protein, other minor polysaccharides
such as heteromannans, and some phenolic acids, chiefly feru-
lic acid (Table 1). Endosperm walls generally lack lignin.
Although cell wall components account for only 510% by
weight of the grain, they can have a disproportionately large
impact on grain technology, utilization, and nutrition.
Biosynthesis of Cell Wall Polysaccharides in Cereals
Given the central role played by cell wall polysaccharides in
cereal processing, there has been considerable interest in the
enzymes that mediate their biosynthesis, particularly in the
developing grain. Cellularization of the starchy endosperm
early in grain development involves atypical growth of cell
walls around individual nuclei in a multinucleate syncytium.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00102-9
 PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing
Table 1 Composition of cell walls in cereal endosperm
Cereal Source of walls Major polysaccharides
Barley Aleurone 71% Arabinoxylan
26% (1,3;1,4)-b-Glucan
2% Cellulose
2% Glucomannan
Starchy endosperm 75% (1,3;1,4)-b-Glucan
20% Arabinoxylan
2% Cellulose
2% Glucomannan
Wheat Aleurone 65% Arabinoxylan
29% (1,3;1,4)-b-Glucan
2% Glucomannan
2% Cellulose
Starchy endosperm 70% Arabinoxylan
20% (1,3;1,4)-b-Glucan
7% Glucomannan
4% Cellulose
Rice Starchy endosperm 27% Arabinoxylan
20% (1,3;1,4)-b-Glucan
28% Cellulose
3% Pectin
15% Xyloglucan/mannans
The fertilized endosperm mother cell divides for
70 h to
form up to 2000 free, individual nuclei in the cytoplasm of
the central cell. Walls subsequently grow between the free
nuclei. After a lag phase, further expansion of the cellular
endosperm occurs through meristematic activity of peripheral
cells that eventually differentiate to form the aleurone. In
barley, the major components of the starchy endosperm
walls, (1,3;1,4)-b-glucan and arabinoxylan, accumulate
steadily between 13 and
40 days post anthesis (dpa).
Despite the importance of cell wall biosynthesis as a deter-
minant of final cereal grain quality, polysaccharide synthases
involved in wall synthesis have proved very difficult to purify
by conventional biochemical procedures, mainly because they
are membrane-bound enzymes and can be rapidly inactivated
in vitro. A key breakthrough in this area occurred when cellu-
lose synthase genes (CesA) were identified in higher plants;
subsequently, multiple CesA genes and multiple groups of
cellulose synthase-like (Cs1) genes have been identified. It
has now been demonstrated that the CslF, CslH, and CslJ
genes encode (1,3;1,4)-b-glucan synthases and that the CslF
group is probably the most important in cereal grain develop-
ment. Indeed, mutations of the CslF6 gene result in grain with
little or no (1,3;1,4)-b-glucan, and when the CslF6 gene is
overexpressed in developing barley grain, (1,3;1,4)-b-glucan
levels increased significantly, as did dietary fiber content of
the grain. Nevertheless, it is becoming increasingly apparent
that other proteins and enzymes are also required for (1,3;1,4)-
b-glucan synthesis and regulation.
Identification of the xylan synthase genes in cereals has
been particularly challenging, and although genes encoding
enzymes from the glycosyltransferase (GT) GT43 and GT47
families have been implicated in the synthesis of the (1,4)-b-
xylan backbone of cereal grain arabinoxylans, precise details of
the roles of these enzymes are now yet available. Enzymes
encoded by the GT61 family appear to mediate the addition
139
of arabinofuranosyl residues to the xylan backbone. There have
been suggestions that xylan synthesis might also involve
enzymes of the cellulose synthase or cellulose synthase-like
enzyme families, given the structural similarities between cel-
lulose and the xylan backbone of arabinoxylans. As the
genomes of more cereal species are sequenced, it is clear that
most genes involved in cell wall polysaccharide synthesis are
members of multigene families.
Enzymic Hydrolysis of Wall Polysaccharides
in Germinated Grain
Most of the enzymes involved in (1,3;1,4)-b-glucan and arabi-
noxylan degradation in the endosperm of germinated grain
have been characterized in detail, particularly in barley,
wheat, and maize. In each case, a battery of hydrolytic enzymes
is required to completely depolymerize the polysaccharides.
The enzymes responsible are generally synthesized de novo in
the aleurone layer or the scutellum and subsequently secreted
into the starchy endosperm. Walls of the starchy endosperm
are completely degraded during this process, but a thin inner
layer of the aleurone cell walls appears to resist degradation,
presumably to maintain the structural integrity of the aleurone
cells while enzyme secretion is in progress.
The (1,3;1,4)-b-glucans of endosperm cell walls are depo-
lymerized by (1,3;1,4)-b-glucan endohydrolases, of which
there are one or two isoforms in most cereal species. The
(1,3;1,4)-b-glucan endohydrolases release low-molecular-
mass (1,3;1,4)-b-oligoglucoside products that can be further
hydrolyzed by exohydrolases such as b-glucan glucohydrolases
and b-glucosidases. The gene families that encode these
enzymes are generally larger, with five or more members.
A second, distinct class of (1,3;1,4)-b-glucan endohydrolase
has been reported in germinated barley and in maize coleop-
tiles. The enzymes release (1,3;1,4)-b-glucans with degrees of
polymerization of 60100, but the enzymes have not been
characterized in detail. Their action patterns are consistent
with a cellulase, but there remains some doubt as to their
precise role in (1,3;1,4)-b-glucan depolymerization in germi-
nated grain.
Arabinoxylan depolymerization in germinated cereal grains
also requires the concerted action of a range of different hydro-
lytic enzymes. (1,4)-b-Xylan endohydrolases hydrolyze glyco-
sidic linkages in the (1,4)-b-xylan backbone of the
polysaccharide and probably require a short, unsubstituted
region of the (1,4)-b-xylan chain for activity. Genes and
cDNAs encoding barley (1,4)-b-xylan endohydrolases have
been isolated and used to define the expression patterns and
chromosome locations of the genes. There is another class of
enzymes that can hydrolyze polymeric arabinoxylans from
cereal cell walls. These enzymes hydrolytically remove single
a-arabinofuranosyl substituents from the (1,4)-b-xylan back-
bone of the polysaccharide and have been designated arabi-
noxylan arabinofuranohydrolases (AXAH). They are probably
responsible for the removal of arabinofuranosyl residues prior
to (1,4)-b-xylan endohydrolase action in germinated grain but
might also be involved in the modification of arabinoxylan
fine structure during wall deposition, maturation, or expan-
sion. Despite these possibilities, there has not been any defin-
itive description of the role of the AXAHs in cereal grain
140 PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing
development or germination. Complete depolymerization of
oligoarabinoxylosides released from cell wall arabinoxylans by
(1,4)-b-xylan endohydrolase requires the action of a-
arabinofuranosidases and b-xylosidases, both of which have
been detected in germinated cereal grains. Enzymes with dual a-
arabinofuranosidase/b-xylosidase activity have been described
and again these hydrolases are encoded by multigene families.
Enzymes in Grains That Affect Processing, End-Product
Appearance, Nutritional Value, or Storage of Grain
This section will focus primarily on wheat that is a staple food
in many parts of the world and unique in terms of the myriad
of end products that are consumed.
a-Amylases in Developing and Ripe Wheat Grains
Starch constitutes up to 80% of the material in the endosperm of
cereal grains and is stored in a highly crystalline form in granules
that are laid down during development and ripening. a-Amylase
is a critical enzyme in remobilization of the endosperm since it
initiates hydrolysis of the starch granules, reducing the large,
complex molecules of starch to smaller units as already
described. Wheat and barley grains can synthesize two groups
of a-amylases, or a-amylase isoenzymes, that differ in genetic
control, temporal pattern of synthesis, and physical and chem-
ical properties. Perhaps, the most important group in grains are
the high-pI (germination or malt) a-amylases that are products
of the a-Amy1 genes that are normally only synthesized in the
grains during the early stages of germination. The second group,
low-pI (pericarp or green) a-amylases, are products of the a-
Amy2 genes. They are present in the pericarp of immature grains,
at low levels in mature grains, and are also synthesized during
the later stages of germination. In comparison with other cereal
enzymes, the a-amylases are relatively thermostable and remain
active at temperatures above 60  C where the starch is gelati-
nized and becomes extremely sensitive to enzymic hydrolysis.
Under ideal growth conditions, ripe wheat grains contain a
small amount of low-pI a-amylase and no high-pI a-amylase,
and following milling, the flour can be used to produce a vast
array of processed foods including pan breads, flat breads,
noodles, cakes, and biscuits as well as industrial products. For
leavened products such as western-style pan breads, some a-
amylase is required to produce the fermentable sugars needed
by the yeast. This a-amylase, often microbial in origin, is
normally added in measured amount during mixing particu-
larly in the highly automated bread-making plants that require
strict adherence to specifications to maintain quality and effi-
cient production. Under nonideal growing conditions, or as a
result of adverse genotype  environment interactions, variable
amounts of either, or both, groups of a-amylases may be
retained or synthesized prior to harvest and as a consequence
the grain may be unsuitable for processing. Wheat with unac-
ceptably high levels of a-amylase is commonly downgraded in
quality, often to be restricted to animal feed, and attracts
substantial financial dockages. While some of the a-amylase
is removed with the bran and germ during milling, at progres-
sively higher levels of activity, more of the enzyme will have
been transported into the endosperm and will be recovered in
the flour. Excess levels of a-amylase in wheat have a disastrous
effect in long fermentation processes, which applies to most
types of western-style pan breads and leads to sticky dough that
is harder to handle, a sticky crumb, dark-colored crust and
loaves with large holes that are difficult to slice and may jam
slicing machines. During cooking or baking, the starch
becomes increasingly susceptible to attack by a-amylase, and
even small amounts of enzyme can cause dramatic differences
in starch viscosity and product texture. Flat breads, chapatis,
and sourdough breads are more tolerant to a-amylase, whereas
Japanese-style (udon) noodles are very sensitive. Irrespective
of the end product, appearance and quality deteriorate as a-
amylase activity increases.
High levels of a-amylase activity in ripe wheat grain can
arise via a number of mechanisms. First, the low-pI a-amylase
that is synthesized in the green (chlorophyllous) pericarp of
immature grains may be retained rather than being gradually
degraded as the grain matures (Figure 1).
Frost damage to partially developed grains and conditions
of high humidity and low light intensity during ripening have
been associated with this abnormal retention of low-pI a-amy-
lase. Second, some bread and durum wheat genotypes contain
a genetic defect referred to as late-maturity a-amylase (LMA) or
prematurity a-amylase (PMAA) that may be result in the syn-
thesis of high-pI a-amylase in the aleurone tissue during the
mid to later stages of grain development (Figure 1). Induction
of high-pI a-amylase synthesis in LMA-prone varieties may be
constitutive, in tall wheat varieties and in semidwarf varieties
grown under mild conditions free from moisture stress, or
require a cool temperature shock in semidwarf varieties ripen-
ing under warm to hot conditions. LMA, unlike germination or
sprouting, is not dependent on rain and does not appear to
involve changes in, or signals from, the embryo. Finally,
adverse weather conditions (rain accompanied by high humid-
ity) at maturity and prior to harvest may trigger grains to
commence germination while still in the spike. This phenom-
enon, known as preharvest sprouting (Figure 2), is relatively
common in many parts of the world.
Preharvest sprouting-resistant varieties in which the grain is
still dormant at harvest ripeness have been developed to combat
this problem. The sequence of events that occurs during sprout-
ing follows very closely those observed in isolated grain that is
germinated under more controlled conditions. a-Amylase syn-
thesis begins adjacent to the embryo or germ end of the grain
and progresses in a wave toward the distal end. The system has
been very widely studied by plant biochemists endeavoring to
understand the intricacies of control of enzyme synthesis and
the role of plant hormones. Initially, only high-pI isoenzymes
are synthesized; however, as germination progresses, the low-pI
groups are also produced. While LMA and sprouting are readily
distinguished both in genetic control and in the pattern of
enzyme distribution within the grain, the a-amylase isoenzymes
involved are similar, as are the effects on the quality of end
products. In contrast to LMA, where only high-pI a-amylase
appears to be synthesized, sprouted grains may contain elevated
levels of a range of hydrolytic enzymes including endoproteases,
lipases, and cell wall-degrading enzymes.
Endoproteases in Developing and Ripe Wheat Grains
Endoproteases attack the internal peptide bonds of proteins
and may make them more accessible to exoproteases that
 PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing 141

50

45

40

35
-Amylase activity

30

25

20

15

10

0
0 5 10 15 20 25 30 35 40 45 50
Days after anthesis
Sprouted Nonsprouted
70 Figure 2 Sprouted (left) and nonsprouted (right) spikes of wheat.

60
critical to the production of high-quality end products such as
bread and noodles.
50
-Amylase activity

40 Enzymes Involved in Determining the Color and Appearance

30
20
10
0 color stability, because these characteristics strongly influence
0 24 48
Time (h)
Figure 1 Diagrammatic representation of a-amylase synthesis in wheat
grains during development, ripening, and preharvest sprouting. Low-pI
a-amylase is synthesized in the pericarp during the early stages of grain
development and then declines to very low levels at maturity in all
genotypes (a), while new low-pI a-amylase is synthesized by the
aleurone later in grain germination or sprouting (b). High-pI a-amylase is
synthesized in the aleurone during later stages of grain development in
LMA-prone cultivars (a) and in the aleurone during the early stages of
germination of all cultivars (b). Low-pI a-amylase (thick line), high-pI
a-amylase (thin line).
of End Products such as Asian Noodles
Large quantities of the wheat used in Southeast Asia, Japan,
Korea, and China are consumed as noodles and increasingly
these products are also finding a place on tables in western
societies. As the markets for noodle products develop and
become more quality-sensitive, greater emphasis has been
placed on improving appearance, consistency of color, and
customer appeal. Asian noodles made from wheat flour come
in a wide range of forms, although there are two basic recipes
used for production. These are referred to as white salted
noodles (WSN), made from flour, water, and varying amounts
of salt, and yellow alkaline noodles (YAN), prepared using
flour, water, and alkaline salts such as potassium and sodium
carbonate. For both types, initial color should be clean and
bright and there should be minimal darkening in the time
between preparation and cooking. WSN vary from white to
cream in color, while YAN have a yellow color that is often
enhanced with artificial coloring agents or eggs.

Synthesis of Flavonoids and Xanthophylls

cleave amino acids from the amino or carboxyl terminals of
peptides and proteins. There are at least four groups of endo-
proteases in wheat grains that differ in their tissue location,
their temporal control during grain development or germina-
tion, and their substrate specificity. Three of these are present
in various tissues in developing grains, but it is not clear
whether they can degrade the major grain storage proteins. In
contrast, acidic thiol endoproteases that are low in developing
grain are synthesized during germination. These endoproteases
can attack the gliadin and glutenin proteins responsible for the
unique rheological and processing properties of wheat that are
The creamy color of WSN and a part of the yellow color of YAN
are dependent on natural yellow pigments, xanthophylls (pri-
marily lutein), present in the germ and endosperm of wheat
grains. Lutein is a member of the carotenoid family of com-
pounds that are found in all plants and are synthesized from
isopentenyl phosphate via a biosynthetic pathway commenc-
ing with geranylgeranyl pyrophosphate synthase. The addi-
tional yellow color that develops in the presence of alkaline
salts in YAN is attributed to flavonoid compounds such as
flavone glycosides that are found in the germ and seed coat
but not the endosperm, as well as unidentified chromophores
that appear to be located in the endosperm and associated with
142 PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing
protein. The starting point for synthesis of flavonoids is phe-
nylalanine and the enzyme phenylalanine ammonia lyase,
which leads into a biosynthetic pathway that produces a vast
array of structures including flavones, cinnamic acids and lig-
nins, anthocyanins, and tannins. Xanthophylls and flavonoids
accumulate during the early to middle stages of grain develop-
ment, depending on the activity of the enzymes in the respec-
tive biosynthetic pathways. In the case of xanthophylls, the
levels decline markedly as the grain ripens. Lutein may also
be converted to mono- and di-fatty acid esters during storage of
ripe grain or flour. The concentrations of xanthophylls and
flavonoids in flour are a product not only of the balance of
synthesis and degradation during grain development but also
of the milling process, in particular the efficiency of separation
of starchy endosperm from other grain tissues such as the germ
and seed coat.
Lipoxygenase
Lipoxygenases (LOX) catalyze the oxidation of polyunsatu-
rated fatty acids such as linoleic and linolenic acid to hydro-
peroxides. Fatty acid radicals produced during the intermediate
steps in the reaction can initiate oxidative degradation of pig-
ments such as lutein and b-carotene. As a result of this co-
oxidation, lutein and lutein ester pigments may decline
during dough mixing in preparation for the production of
WSN and durum pasta, leading to poor color stability. Loss
of pigment can be reduced by the addition of compounds such
as a-tocopherol that inhibit LOX activity or by using bread and
durum cultivars that have been selected for zero LOX activity.
In addition to its effects on color, LOX has been associated with
the formation of compounds with undesirable odors and fla-
vors in a range of products. Loss of lutein and carotene is also
undesirable due to the role these compounds play in the health
of our eyes, in iron uptake in the intestinal tract, and as general
scavengers of peroxy radicals that reduce oxidative damage to
tissues.
Polyphenol Oxidase
Polyphenol oxidases (PPO) catalyze the oxidation of phenolic
compounds to produce brown pigments on the cut or dam-
aged surfaces of fruits and vegetables and can cause major
economic losses. PPO located in the seed coat of ripe wheat
grains is able to hydroxylate certain phenols in the s-position
adjacent to the existing OH group to form orthodiphenols.
Figure 3 Yellow alkaline noodle sheets: from left to right control (2% sodium carbonate), 10 mM tyrosine (polyphenol oxidase substrate),
and 10 mM tyrosine plus 10 mM tropolone (specific polyphenol oxidase inhibitor).
The orthodiphenols are oxidized further by PPO to s-benzo-
quinones, which in turn polymerize nonenzymatically to mel-
anins (brown pigments). Fortunately, most of the PPO in
wheat grains is removed with the bran during milling, but
sufficient enzyme is carried through in the flour of many
wheat cultivars to present problems with the color stability of
products that are not cooked immediately. While the endoge-
nous substrate of PPO in grains has not been identified, the
role of PPO in darkening of WSN, YAN, and durum pasta is
now clear. Addition of exogenous substrates such as tyrosine,
catechol, and L-DOPA (L-3,4-dihydroxyphenylalanine)
enhances noodle darkening, whereas darkening is greatly
reduced in the presence of specific PPO inhibitors or in culti-
vars selected for low or zero grain PPO activity (Figure 3).
Variation in PPO activity explains most of the genetic differ-
ence in noodle darkening between cultivars, but is only respon-
sible for a portion of the total darkening that occurs. Physical
effects and other unidentified oxidation reactions also contrib-
ute to the reduction in noodle brightness over time.
Peroxidases and Hydrogen Peroxide Synthesis and Catalysis
Peroxidases are oxidoreductases widely distributed in higher
plants and have been shown to oxidize many compounds,
including phenolics, in the presence of hydrogen peroxide.
Oxidation of phenolic compounds by these enzymes can lead
to the formation of brown pigments similar to those formed by
PPO, and consequently, they have also been implicated in
browning or darkening of noodles and pasta. In the case of
peroxidase, the situation is complicated by the presence of a
number of forms with different substrate specificities that are
localized in embryo, endosperm, and seed coat tissues of wheat
grains and by the interaction with systems that either generate
H2O2 (superoxide dismutase and NADH oxidation) or catalyze
(catalase) its destruction.
In addition to its possible role in product darkening, per-
oxidase has been implicated in the development of symptoms
of a grain quality defect known as black point. Both wheat
and barley are prone to this defect, which manifests as a dark
stain in the seed coat or husk, respectively, at the embryo end
of the grain and is associated with warm, humid conditions
during ripening (Figure 4). Unfortunately, this pigmented
tissue fragments during milling and black specks are carried
through into wheat flour or semolina. While it has no effect on
processing properties or texture, the specks are visible in
 PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing
Figure 4 Sound and black point affected wheat grains.
noodle and pasta products and high levels of black point result
in downgrading and substantial economic losses.
The enzymes discussed in this section all have specific
functions within the tissues of developing grains, although in
some instances the exact role has not been identified unequiv-
ocally. Their physiological functions are quite distinct from
their effects on processing and end-product appearance.
While in many cases it is possible, or at least conceivable, to
develop germplasm with extremes of variability for levels of
these enzymes, some caution is required in case such dramatic
changes have adverse effects on plant growth, disease resis-
tance, or grain yield.
Enzymes that Are Active in Ripe, Quiescent Grains
Despite the low moisture content of ripe cereal grains and the
death of the seed coat and starchy endosperm tissues during
grain ripening, it is clear that respiration and some other
enzyme-catalyzed reactions continue at a slow rate during stor-
age of harvested grain. Some of these activities result in the
modification of the structure of grain constituents and secondary
metabolites, while others may impact on the storage quality or
the capacity to maintain freshness and palatability. In addition,
ripe grains contain a range of enzymes such as exopeptidases and
b-amylase that are sequestered in active or inactive form. These
enzymes can spring into action once endo-acting enzymes,
synthesized during germination or sprouting, catalyze the initial
depolymerization of protein and starch.
At maturity and depending on the genotype and growing
conditions, wheat grains contain varying levels of xanthophyll
(hydroxy carotene) present primarily as free lutein. During
storage of wheat grain and wheat flour, and again depending
on genotype as well as temperature and humidity, considerable
proportions of the free lutein (up to 60%) may be converted to
mono- and di-fatty acid esters. A specific GDSL-lipase (the
prefix GDSL is added to distinguish this group of lipases from
common lipases and refers to specific sequence of amino acids
at the active site of this group of lipases), coded by a gene
located on chromosome 7D, appears to be responsible for
esterification possibly acting as a trans-esterase to transfer lino-
leic and palmitic acids from membrane phospholipids to the
hydroxyl groups of lutein. The lipase is synthesized during the
early to middle stages of grain development but does not
catalyze esterification of lutein until the ripe grain is stored
under warm, dry conditions. Esterification does not occur in
143
durum wheat that lack the D-genome Wheat Genomics or in a
small number of bread wheat cultivars that contain a non-
functional version of the gene. Since adequate lutein intake is
associated with a reduction in the incidence of glaucoma and
macular degeneration, a major cause of blindness in elderly
people, this modification may have implications for absorp-
tion, bioavailability, and attempts to improve eye health
through dietary supplementation.
Enzymes have been implicated in the deterioration in qual-
ity of rice during storage. This deterioration may be linked to
lipid peroxidation, peroxidative changes in polyunsaturated
fatty acids that in turn are governed by the balance of produc-
tion and dissipation of reactive oxygen species. Peroxidases are
key scavengers of reactive oxygen species and recent studies
have indicated that the activity of this enzyme declines during
storage of rice.
Endogenous Enzyme Inhibitors
Cereal and legume grains contain inhibitors against a range
of enzymes such as proteinase, a-amylase, endoxylanase, and
limit dextrinase. While some inhibitors are only effective
against exogenous enzymes of microbial or insect origin,
others are capable of inhibiting endogenous grain enzymes.
The physiological function of these inhibitors is not certain but
it has been suggested that they may be involved in plant
defense mechanisms, protecting the stored reserves in the
seed from invading microbes and insects, and/or play a regu-
latory role in plant development and metabolism. A role in the
plant defense mechanisms seems plausible given the effective-
ness of many endogenous inhibitors against hydrolytic
enzymes from microbial and insect sources. Support for a
regulatory role is less plausible and requires unequivocal evi-
dence that the inhibitors reduce the activity of particular
endogenous grain enzymes at physiological concentrations
and that the inhibitors are located in the same tissue as the
target enzyme. Most of the proteinase inhibitors that have been
characterized to date are inhibitors of a particular class of
protein hydrolases, the serine proteinases. As yet, there has
been no conclusive demonstration of serine proteinases of
the chymotrypsin family or their genes in plants. In the case
of the bifunctional a-amylase/subtilisin inhibitor that is com-
mon in barley grains, there is clear evidence that it can signif-
icantly reduce the activity of endogenous high-pI a-amylase
extracted from germinated or malted grain and ameliorate
144 PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing
some of the deleterious effects of high-pI a-amylase on loaf
volume and structure. However, recent studies of gene expres-
sion have indicated that the inhibitor is primarily located in
the maternal seed coat tissues of barley grains rather than
endosperm. Similarly, whereas proteins of the serpin super-
family are localized in both vegetative and endosperm tissues
of barley grains, they only inhibit specific mammalian serine
proteinases. Limit dextrinase inhibitors isolated from barley
inhibit the activity of endogenous limit dextrinase and appear
to be ineffective against a number of other enzymes capable of
hydrolyzing a-1,6-linkages in dextrins. In theory at least, these
inhibitors could interfere with the malting process although
a role in commercial malting has not been demonstrated.
Two groups of endoxylanase inhibitors (XIP-type and TAXI-
type) have been isolated from cereal grains that are effective
against fungal endoxylanases and both bacterial and fungal
endoxylanases, respectively. Microbial endoxylanases are
commonly used in bread making to improve loaf volume.
Irrespective of possible roles in plant defense and regula-
tion, significant numbers of inhibitors are active either against
endogenous enzymes that are important in processing or
against exogenous enzymes, usually of fungal or bacterial ori-
gin, that are added as ingredients to improve processing and
quality of end products. In these instances, the inhibitor and
the enzyme are brought into contact as a result of milling or
deliberate physical admixture and there may be important
implications for industrial applications.
Future Prospects
Many cereal products and processes depend on, or are
adversely affected by, specific enzymes either that are present
in grains at maturity or that develop as the result of interactions
between grains and the environment. As more of these
enzymes are recognized, and their impact quantified, many
opportunities will be presented to improve products and to
develop new and novel products and processes. In many cases,
there is already sufficient genetic variation within existing
germplasm or near relatives for the selection of lines with
enzyme levels more appropriate to the particular end-use pro-
cess, while in other instances, it may be possible via mutation,
genetic engineering, or modification of milling techniques to
extend variation beyond that currently available. Genetic engi-
neering, in particular, opens up the possibility of introducing
genes encoding novel enzymes into cereal grains, overexpres-
sing genes controlling synthesis of existing enzymes, or synthe-
sizing enzymes in tissues such as the starchy endosperm where
they are currently absent. As an example, identification of the
genes should reveal potential strategies for the manipulation of
cell wall (1 ! 3,1 ! 4)-b-D-glucan and arabinoxylan levels, for
the enhancement of cereal quality and productivity. The 3-D
structure of one of the barley (1,3;1,4)-b-glucan endohydro-
lases has been solved and the structural information used to
rationally design mutated forms of the enzyme with enhanced
thermostability. The increased thermostability of this barley
(1,3;1,4)-b-glucan endohydrolase could have commercial
potential through its ability to overcome filtration problems
and other difficulties encountered in the malting and brewing
industries. Increasingly sophisticated and high-throughput
genomics technologies, coupled with rapid functional analysis
systems, are now available to researchers. These new technol-
ogies can be confidently expected to identify more genes and
reveal potential strategies for further manipulation of enzyme
and grain constituent levels relevant to the grains industries in
the near future.
Exercises for Revision
What enzymes are important in the development and
breakdown of endosperm cell walls?
What enzymes are important during germination?
When are the different a-amylase families synthesized in
wheat grains?
Which enzymes present in ripe wheat grains affect end-
product quality?
What methods are used to overcome the deleterious effects
of some enzymes on the quality of wheat-based end
products?
What is black point and does it affect end-product quality?
Exercises for Readers to Explore the Topic Further
Fully characterize all the gene families involved in endo-
sperm (aleurone and starchy endosperm) cell wall synthesis
and degradation, develop a better understanding of their
respective roles and interactions, and explore ways of mod-
ifying their activity to improve products and processing.
Genetic and molecular characterization of late-maturity a-
amylase in wheat to assist in the development of varieties
that maintain low a-amylase activity and to provide new
insights into the processes involved in the control of grain
development.
Utilize genetic and molecular techniques to optimize the
grain content of enzymes and constituents that impact on
end-product quality and grain storage without compromis-
ing productivity.
See also: Barley, Rice and Maize Processing: Barley: Malting;
Carbohydrates: Cereals: Chemistry and Physicochemistry of Non-
starchy Polysaccharides; Starch: Synthesis; Fats: Lipid Chemistry;
Genetics of Grains: Wheat Genetics and Genomics; Proteins:
Protein Synthesis and Deposition; The Basics: Grain: Morphology of
Internal Structure; The Cereal Grains: An Overview of the Family of
Cereal Grains Prominent in World Agriculture; Wheat Processing:
The Gluten Proteins of the Wheat Grain in Relation to Flour Quality;
Wheat-Based Foods: Noodles: Asian Wheat Flour Noodles.
Further Reading
Asenstorfer RE, Appelbee MJ, and Mares DJ (2009) Physical-chemical analysis of non-
polyphenol oxidase (non-PPO) darkening in yellow alkaline noodles. Journal of
Agricultural and Food Chemistry 57: 55565562.
Chen TF and Chen CL (2003) Analysing the freshness of intact rice grains by colour
determination of peroxidase activity. Journal of the Science of Food and Agriculture
83: 12141218.
Dominguez F and Cejudo FJ (1996) Characterization of the endoproteases appearing
during wheat grain development. Plant Physiology 112: 12111217.
 PROTEINS | The Enzymes Associated with Food Grains and Their Functions in Processing
Fincher GB (1989) Molecular and cellular biology associated with endosperm
mobilization in germinating cereal grains. Annual Review of Plant Physiology and
Plant Molecular Biology 40: 305346.
Fincher GB and Stone BA (1986) Cell walls and their components in cereal grain
technology. Advances in Cereal Science and Technology 8: 207295.
Fuerst EP, Anderson JV, and Morris CF (2006) Delineating the role of polyphenol
oxidase in the darkening of alkaline wheat noodles. Journal of Agricultural and Food
Chemistry 54: 23782384.
Garcia-Olmedo F, Salcedo G, Sanchez-Monge R, Royo J, and Carbonero P (1987) Plant
proteinaceous inhibitors of proteinases and a-amylases. Oxford Surveys of Plant
Molecular and Cell Biology 4: 275334.
Hidalgo A and Brandolini A (2012) Lipoxygenase activity in wholemeal flours from
Triticum monococcum, Triticum turgidum and Triticum aestivum. Food Chemistry
131: 14991503.
145
Howitt C and Pogson BJ (2006) Carotenoid accumulation and function in seeds and
non-green tissues. Plant, Cell and Environment 29: 435445.
Mares D and Mrva K (2008) Late maturity a-amylase: low falling number in the absence
of preharvest sprouting. Journal of Cereal Science 47: 617.
Shirley BW (1998) Flavonoids in seeds and grains: Physiological function,
agronomic importance and the genetics of biosynthesis. Seed Science Research
8: 415422.
Stewart RJ, Varghese JN, Garrett TPJ, Hoj PB, and Fincher GB (2001) Mutant barley
(1 ! 3,1 ! 4)-b-glucan endohydrolases with enhanced thermostability. Protein
Engineering 14: 245253.
Williamson PW (1997) Black point of wheat: In vitro production of symptoms, enzymes
involved and association with Alternaria alternate. Australian Journal of Agricultural
Research 48: 1319.
 Proteomic Analysis Relevant to Cereal Grains
A Juhasz, Centre for Agricultural Research of the Hungarian Academy of Sciences, Martonvasar, Hungary
R Haraszi, Campden BRI, Gloucestershire, UK
F Bekes, FBFD PTY LTD, Beecroft, NSW, Australia
DJ Skylas, Grain Growers Limited, North Ryde, NSW, Australia
CW Wrigley, The University of Queensland, Brisbane, QLD, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Definition of the concept of proteomics versus other types
of -omics.
Display of the proteome as a two-dimensional map of
spots, using gel-based technology.
Gel-free proteomic analysis and LC fractionation.
Qualitative and quantitative identification of the
polypeptides.
Mass spectrometry-based proteomics.
Proteome mining and functional proteomics.
Learning Objectives
To achieve an understanding of proteomics and its relation-
ships to genomics and to the functions of individual cereal
grain polypeptides.
To understand how proteomics can provide greater insights
into the physiological functions of the cereal grain and into
the utilization of grains.
Introduction
Elucidation of grain composition at the molecular level is
needed to permit ongoing genetic improvements to grains
with respect to their efficient production and processing. This
information is especially needed concerning the proteins of the
grain, because they are the primary molecules in implementing
the directions from the relevant genes, under the further
control of environmental conditions. There is thus an impor-
tant interaction between genotype (the variety, comprising the
set of genes) and growth conditions (Figure 1), and this inter-
action determines the synthesis of the individual proteins that
are needed to conduct all the functions of the plant cells.
Background to Proteomics
The study of proteins and their functions has long been an
accent of biochemical research, but it has been only in the
1990s that the concept of proteomics has been introduced
to this study. Proteomics is a powerful approach to the analysis
of gene expression in biological systems, with the term
proteome indicating the full complement of proteins
expressed by the genome of an organism (e.g., wheat) in a
tissue (e.g., leaf, endosperm, or embryo) at a particular stage
of development and under specific growth conditions. Even
146 Encyclopedia of Food Grains, Second Edition
though there is only one definitive genome of an organism, it
can code for multiple expressed proteomes. The expression of a
single protein largely depends on the environment and is a
result of combined events including transcription, translation,
and protein turnover and posttranslational modifications
(PTMs). The specific genes that are active are also determined
by the growth conditions, as shown in Figure 1.
The first activity of the genes is transcription, that is, the
production of messenger ribonucleic acid (mRNA), with a
nucleotide sequence based on that of the stretch of DNA that
constitutes the particular gene. Analysis of the mRNA (the
transcriptome) provides information about which proteins
are to be synthesized in the next stage of translation, as
shown in Figure 1. Routinely, analysis of the proteins (the
proteome) is preferred over study of the mRNA, because the
mRNA molecules are transitory, being degraded after their
role in protein synthesis is completed. Therefore, the mRNA
transcripts may not be a true indicator of protein expression
and often do not correlate with protein levels.
The transcriptome and the proteome are thus subsets of the
wider range of mRNA and protein molecules that might have
been produced, given the full extent of the genome. In sum-
mary, the study of the proteome tells us about the range of
proteins that relate to the specific tissue under study (root or
shoot or mature grain), as it was grown under the particular
environmental conditions existing at the time for the genotype
involved.
Methods of Proteomic Analysis
Sample Preparation
Seed tissues, such as bran and seed endosperm, contain a large
number of diverse proteins, with different chemical character-
istics that resulted in the development of special extraction and
purification requirements. To extract seed proteins in a satis-
factory amount and purity, there are several basic criteria that
need to be considered. For cereal grains, the major storage
components are starch and different carbohydrates that have
an adverse effect on the extractability of proteins. Compared to
other plant tissues, extraction of seed proteins first requires the
mechanical breakage of the hard structure of the mature seed
using grinding or milling. This can be followed by the subse-
quent solubilization-based Osborne fractionation or the use of
either proteolytic enzymes or special chemicals in order to
further break down or fractionate polypeptides. Purification
and protein enrichment can be essential steps, too, which
also require the use of sophisticated and expensive chemicals.
Due to frequent interference between chemicals and sensitive
http://dx.doi.org/10.1016/B978-0-12-394437-5.00107-8
 PROTEINS | Proteomic Analysis Relevant to Cereal Grains 147

Genomics Transcriptomics Proteomics Metabolomics Discovery

DNA RNA Proteins Modified proteins Biological function

Gene
expression Translation Phenotype
Post-translational
modifications

Metabolites
Sugars, nucleotides, amino acids, lipids

Environment

Figure 1 Newly emerging technologies to elucidate genome and proteome interactions.

Figure 2 Workflow of the gel-based and gel-free methods of proteomic analysis.

detection techniques (e.g., NaCl ionization in the ESI of a mass
spectrometer suppresses the peptide ionization), they need to
be removed at a proper time of analysis. Starch and complex
carbohydrates can also seriously reduce protein extraction,
which might be a primary problem in proteomic analyses of
maize or rice seeds, however. The ideal extraction method
should be compatible with downstream protein analysis by
gel electrophoresis, liquid chromatography, and mass spec-
trometry and subsequent protein identification (Figure 2).
Gel-Based Methods
Gel-based proteomics is still the main approach used in grain
proteomics. For decades, one-dimensional gel electrophoresis
148 PROTEINS | Proteomic Analysis Relevant to Cereal Grains
(1-D GE) was the most popular method in protein chemistry
due to its ability to determine and compare the molecular
weights of proteins. It was used in different approaches, such
as to compare and determine the number and size of seed
storage proteins in wheat. It is also used in immunodetection
or Western blotting. Today, it is most often used as the second
dimension in two-dimensional gel electrophoresis. Two-
dimensional gel electrophoresis (2-D GE), introduced in
1975, is the most widely used gel-based technique to separate
proteins. The first dimension involves isoelectric focusing
(IEF), in which proteins are fractionated based on their isoelec-
tric points (pI) across a specific pH range using commercially
available immobilized pH gradient (IPG) gel strips. The
second-dimension of fractionation resolves the proteins on
the basis of their molecular mass, using sodium dodecyl sulfate
polyacrylamide gel electrophoresis (SDS-PAGE). The SDS
detergent binds to the denatured proteins, overriding any
intrinsic charge differences, to provide a uniform charge-to-
mass ratio, allowing proteins to resolve according to their
respective sizes. Due to its excellent resolving power, the tech-
nique is suitable to separate over 1000 proteins on a single gel.
The 2-D GE proteomic map in Figure 3 represents the pro-
teins of immature wheat grain flour. It shows the polypeptides
extracted from the immature endosperm of the soft wheat
variety Wyuna, the endosperm being the floury part of the
grain, excluding the germ, embryo, and bran layers. The pro-
teome map of Wyuna appears in two parts, reflecting two
different pH ranges used in IEF. Protein extracts were fraction-
ated in both acidic (pH 47) and basic ranges (pH 611). The
proteomic pattern is thus a map of pI, in the horizontal dimen-
sion increasing from left to right, and of molecular size, in the
vertical dimension. SDS-PAGE provides fractionation on the
general basis of the size of the protein molecules, from largest
(top) to smallest (bottom).
After separation, the proteins in the gels must be visualized.
There are various stains available for protein visualization,
Wyuna cultivar-endosperm proteome maps
4 pH
1
2 6
Figure 3 The acidic and basic proteome maps of immature (17 days postanthesis) wheat grain endosperm of the soft Wyuna variety. Protein spots
were characterized by N-terminal amino acid sequencing and identified from protein-sequence database interrogation.
including Coomassie or silver staining, but the more common
and routinely used stains now are fluorescent dyes, such as
SYPRO Ruby protein stain or CyDyes (Cy2, Cy3, and Cy5).
Two-dimensional Fluorescence Difference Gel Electrophoresis
(2-D DIGE) was developed to overcome problems related to
moderate gel-to-gel reproducibility observed in 2-D gel electro-
phoresis. In this approach, two protein samples and an internal
standard are labeled prior the gel electrophoresis with one of
the three size- and charge-matched, spectrally resolvable fluo-
rescent dyes. The three samples are then mixed and uploaded
onto a single IPG strip followed by SDS-PAGE. The simul-
taneous separation of multiple fluorescent-labeled samples,
together with an internal standard, results in fewer gels needing
to be analyzed in a significantly shorter time frame. Using the
internal standard, the gel-to-gel variation can be eliminated as
no technical replicates are needed to confirm differences in
protein abundance.
Gel-Free Methods: LC-MS and LC-MS/MS Methods
In proteomics, high-throughput protein identification is usu-
ally done by mass spectrometry (MS) or coupling it with more
sensitive methods. Shotgun or bottom-up proteomic
approaches are targeting the analysis of a peptide mixture
that is a derivative of the target proteome. Proteolytic enzymes
or specific chemicals can be used to digest proteins before
chromatographic separation that is followed by peptide ioni-
zation and detection. The general use of trypsin digestion failed
in the case of gluten, which contains low level of arginine and
lysine. Replacement by chymotrypsin was first shown to effec-
tively produce peptides suitable for chromatography. A novel
combination of direct enzymatic digestionLC-MS/MS (ESI
ion trap) method was developed for the semiquantification
of gluten traces in food samples, which applies a two-step
multienzymatic digestion. Recent studies are more and more
acknowledging the odd nature of cereal proteins and
76 pH 11
4
5

demonstrating the need to use unconventional proteomic
approaches for proteins like gluten.
Top-down proteomics involves the gas-phase ionization of
intact proteins and subsequent high-resolution mass measure-
ment of intact protein ions followed by their direct fragmenta-
tion inside the mass spectrometer without prior digestion. Top-
down approaches enable a more complete characterization of
protein isoforms and any PTMs than the bottom-up approaches
but are not exempt from limitations (e.g., high purity, HPLC
separation, and high molecular weight (>50 kDa)). New meth-
odological advances with ultra high performance liquid chro-
matography (UHPLC) and mass spectrometers with ion
mobility are opening up new ways for top-down approaches
in the proteomics field.
Matrix-assisted laser desorption/ionization time of flight
(MALDI-TOF) can be used for indirect quantification of cereal
proteins like gliadins even in processed foods and the direct
identification and exact molecular mass determination of cereal
proteins without prior separation. MALDI-MSPMF (peptide
mass fingerprinting) is a combination, which involves enzy-
matic digestion of proteins resulting in peptide fingerprints.
Sensitivity and resolution of LC-MS methods (e.g., MALDI-
TOF and ESI-Q) can be increased with the application of
coupled (tandem MS) techniques like ESI-QqQ and ESI-Q-
TOF, where selected ionized peptides are fragmented before
entering the detector of the mass spectrometer. The monitoring
of specific mass transitions as SRM (single reaction monitor-
ing) or multiple reaction monitoring (MRM) provides a unique
identification and quantification tool for the analysis of com-
plex mixtures including overlapping peptide characteristics.
Identification and Characterization of Protein Gel Spots
This procedure of 2-D GE has been available for some decades,
providing proteomic maps such as in Figure 3. Six spots are
numbered as examples of polypeptide identification. Table 1
shows the results from excising these six spots from these gels
for identification by N-terminal sequencing. As a result of
matching these sequences to databases, the identities could
be determined for five of these six spots.
Table 1 N-terminal amino acid sequence characterization of immature wheat endosperm proteins, taking the six numbered spots in Figure 3 as
examples
Spot N-terminal
no. sequence Gene product Identity (%)
1 XATFTLPD Superoxide dismutase 87.5% in 8 aa
(EC 1.15.1.1)
2 SGPWMCYV a-Amylase inhibitor 100% in 7 aa
WDAI-3
3 MKLIAAYL 60S acidic ribosomal 100% in 8 aa
protein P2 (ribosomal
protein A)
4, 5 NMQVDPSG g-Gliadin 100% in 8 aa
6 DILRSDQP
Protein sequences were used to interrogate SWISS-PROT and TrEMBL databases via FASTA algorithm.
PROTEINS | Proteomic Analysis Relevant to Cereal Grains 149
The relatively novel aspect of proteomic analysis is the
subsequent stages of postseparation microcharacterization of
the individual protein spots. The ability to characterize and
identify proteins in a high-throughput manner is a critical
factor in proteome projects. Two widely used protein-
characterization methods are N-terminal amino acid sequenc-
ing and MS, with the latter being the preferred method of
choice used in proteome projects.
N-terminal amino acid sequencing has been a popular
method for characterizing proteins in the past. The most com-
mon form of this method is the traditional Edman degradation
chemistry, which can be used to sequence proteins from gels,
or it can be carried out on proteins electroblotted from the
proteome gel to a polyvinylidene difluoride membrane. In
either case, Edman chemistry involves the reaction of phenyl
isothiocyanate with the free amino group of the protein; deriv-
atives are cleaved and then converted to phenylthiohydan-
toins, which differ in their amino acid side chain, allowing
them to be characterized by retention time using reversed-
phased high-performance liquid chromatography. This is not
the preferred method in high-throughput large-scale proteome
projects, due to the relatively low number of samples that can
be processed in parallel (generally only three or four samples),
its low sensitivity, and the time and cost for analysis.
A gel spot can be directly ionized by MALDI and detected
with a TOF or can be destained, reduced, enzymatically
digested, and then extracted (in gel digestion) in order to
have a peptide mixture, which then can be analyzed with LC-
MS or LC-MS/MS for protein composition.
Peptides and proteins are identified from the recorded and
processed mass spectra with a peptide matching algorithm and
against a protein database. Besides experimental conditions
including the chromatographic separation, the precision of
peptide identification is highly depending on the quality of
the mass spectra and the database.
Peptides with completely different amino acid sequences
and functionality may coelute during LC separation, have
identical or very similar masses, and may fragment similarly.
This is particularly true for proteins belonging to a protein
family like alpha-gliadins and low molecular weight (LMW)
Matched
organism Comments and accession no.
Wheat N-terminal sequence starts at residue 22 of O82571 with residues
121 possibly being a signal sequence or truncated. Also
matches to P93606 and Q96185
Wheat P10846 (fragment). Also matches other a-amylase inhibitors
P01084 and P01085
Wheat Subunits P1 and P2 exist as dimers at the large ribosomal subunit
P05390
Wheat N-terminal sequence starts at residue 20 of P21292 as signal
sequence of precursor is residues 119
No close matches
150 PROTEINS | Proteomic Analysis Relevant to Cereal Grains
glutenin subunits. Additionally, the protein content in a gel
spot is usually so little that without sufficient sensitivity and
resolution, the detection and identification of their peptides
are not possible.
Generally, marker peptides are used for the quantitative
detection of proteins. The first step is to do discovery work in
order to map the most unique and detectable peptides using a
high-resolution mass spectrometer like an ESI-Q-TOF or a
software tool such as Skyline or the ProPepper database,
which provides the starting point in the marker selection pro-
cess. After selecting a potential pool of peptides, the next step is
the experimental optimization of the quantification method
using, for example, an ESI-QqQ and synthetic peptides in order
to tune up the mass spectrometer for the accurate detection of
specific predefined mass transitions of the targeted peptides.
The size of a database due to false-positive hits and the
accuracy of entries (proper annotations) are further critical
factors in the relation of peptide matches to the protein source.
This is relevant in the discovery phase and especially complex
for the seed proteome, where the plant genomes are not fully
sequenced yet. In targeted shotgun proteomics, the design of
an SRM for protein quantification needs to fulfill the criteria
that the amount of peptides and that of proteins are equimolar,
which is not the case for, for example, gluten, where the high
level of homology and the multiple prevalence of sequences
limit the establishment of quantitative relationships between a
detected peptide, its individual protein source, its protein
family, and its plant origin.
The Potential Value of Proteomics to Grain Science
Using proteomic analysis, we can identify the individual pro-
teins that are responsible for specific aspects of the phenotype,
Protein identification
Subproteome isolation
Organelle composition
Proteome
mining
Proteomics
Functional and
expressional
Protein interactions proteomics
Signaling pathways
Metabolic pathways
Biological processes
Environmental responses
Gene silencing
Introduction of genes
Figure 4 Proteomic approaches used in grain science. Adapted from Juhasz J, Moolhuijzen P, Bellgard M, Appels R, and Bekes F (2013) Wheat grain
proteomics for the food industry. In: Toldra F and Nollet LML (eds.) Proteomics in Foods. Principles and Applications. Food Microbiology and Food
Safety, vol. 2, pp. 341379. New York: Springer.
that is, the many different characteristics of the plant and of the
grain produced from it. There are different subdivisions of
proteomics that use different approaches and targets to answer
different questions related to protein identification, expres-
sion, function, interactions, etc. These different types are sum-
marized on Figure 4.
Proteome Mining
The water-soluble albumins and salt-soluble globulins of
wheat grain comprise mainly metabolic and protective
proteins, while at least some of the insoluble structural pro-
teins may be extracted with dilute acid or alkali in the glutenin
fraction. Albumins and globulins, each accounting for  10%
of total flour proteins, are known to be soluble enzymes and
proteins soluble in polar solution, respectively. More than
2000 protein spots can be extracted from flour with salt
solution.
The structure of the wheat grain can be categorized simply
into three distinct fractions, the outer layers, the germ, and the
endosperm. The endosperm plays the most important role
from a human consumption point of view and is the fraction
studied in most detail, identifying a large number of proteins
in the developing grain.
The proteins located in the wheat endosperm determine the
characteristics of elasticity, extensibility, and gas-holding abil-
ity of the wheat dough. Particularly important are the proteins
within the gluten complex, the low- and high-molecular-
weight glutenin subunits, proteins, and the diverse group of
gliadins. This group of proteins shows extensive polymor-
phism across wheat genotypes. Wheat also contains smaller
amounts of other storage proteins such as globulins and
triticins and proteins such as amylase and protease inhibitors
Phosphorylation
Glycosylation
Ubiquitination
Polymerization
Post-
translational
proteomics
Computational
and structural
proteomics
Structure function relations
Protein ligand interactions
Enzyme complex analyses
3D protein modelling
Database and bioinformatic analyses
 PROTEINS | Proteomic Analysis Relevant to Cereal Grains
that may protect against insects and fungi. A range of enzymes
exist in very small amounts.
Amphiphilic proteins are detergent-soluble membrane pro-
teins and together with gluten proteins play an important role
in wheat quality. Some of them, like puroindolines, are lipid-
binding proteins and are strongly linked to dough-foaming
properties and to fine crumb texture.
The proteomic investigation of the wheat amyloplast can be
carried out on whole amyloplasts or purified amyloplast mem-
branes, identifying the total number of 171 and 108 proteins,
respectively. The largest class of proteins with a known func-
tion (24%) was found to be starch synthesis and storage
enzymes. Affinity electrophoresis using glucan (cornstarch or
amylopectin) as affinity substrate has been utilized successfully
in the identification of enzyme-type proteins, especially the
proteins related to starch biosynthetic and hydrolyzing
enzymes and their inhibitors.
Achievements of proteomic research on bran and germ
tissues are described by Juhasz et al.
Functional Proteomics
Functional proteomics aims to characterize how changes in
protein localization, expression, and interaction are affected
in the different developmental stages or by different environ-
mental cues. These processes include germination and seedling
development, tillering, stem elongation, head emergence,
flowering, seed development, and dormancy. Functional pro-
teomics is suitable to determine the dynamics of how proteins
are related into complex networks and how their expression
influenced by coexpressing proteins or other biomolecules
present in the tissue. The approaches used are based on quan-
titative analyses of the proteome, combining approaches such
as analytical chemistry, biochemistry, MS, and bioinformatics.
For instance, large-scale proteomic experiments enable us to
understand the key players and mechanisms involved in grain
filling and seed maturation (i.e., the expression of cereal grain
proteins) in order to understand how environmental stresses
affect wheat quality. These mechanisms determine regulatory
principles of transcriptional activation and protein turnover in
response to abiotic or biotic stresses as well as to differences in
developmental processes in the different cereal species or
cultivars.
Expression Proteomics
Quality attributes of a flour sample are defined by the allelic
composition of gluten proteins (the high molecular weight
(HMW) and low molecular weight (LMW) glutenin subunits
together with gliadins) as well as their relative expression
levels. Proteomic analysis, used to monitor the expression of
gluten proteins during grain development, specially the glute-
nin subunits, provides useful information about polymer
formation and helps in relating protein composition to quality
attributes. Nongluten proteins encompass smaller percentages
of the seed protein content; however, they play a crucial role
in different metabolic processes during grain development,
ripening, and desiccation, and they can have a relevant role
in defense mechanisms against biotic and abiotic stresses.
151
Numerous studies have been carried for the identification
and mapping of wheat proteins. Studies, for example, on ker-
nel responses to heat stress have enabled identification of
many of the enzymes in kernel of wheat. However, the major-
ity of soluble proteins still need to be identified and mapped.
Proteomic research activities have been reviewed describing
applications of proteomics in wheat research: dough quality,
starch properties, puroindoline, chromosomal location of cod-
ing genes, effects of growth environment, insect damage to
wheat grains, and genotype identification. The review of Juhasz
el al. provides a table listing 38 application areas in 113 pub-
lications, using the different techniques of proteomics in both
plant biology and food science.
The development of techniques to improve extraction and
separation and to identify proteins and peptides is facilitating
functional proteomics, together with the analysis of sub-
proteomes from small amounts of starting material, such as
seed. The combination of proteomics with structural and func-
tional analysis is increasingly applied to target subsets of pro-
teins. These next-generation proteomic studies differentiate
between special subproteomes and functional subproteomes
and contribute to understanding the processes controlling
cereal grain development, nutritional and processing
characteristics, and the changes caused by biotic and abiotic
stresses. The importance of abiotic stresses caused by global
climate change has particular significance in the future of
agriculture and food industries. Advances in crop proteomics
provide the basis for interpreting changes in the grain as a
result of climatic factors including photosynthetic stress, air
pollutants, thermal stress including heat and cold, osmotic
stress (drought, salt, and flooding), and metal stress to help
to eliminate or to reduce the negative effects of these factors on
the food production chain
Posttranslational Proteomics
Polypeptides produced during translation often undergo dif-
ferent posttranslational alterations by modifications of amino
acid side chains through the addition of a wide range of
molecules, mostly carbohydrates, lipids, phosphate, nitrate,
or methyl groups. These modifications are covalent processing
events that can modulate the activity, localization, turnover,
and interaction of many seed proteins. The resulted modifica-
tions, such as glycosylation, phosphorylation, methylation,
ubiquitination, and deamidation, of the proteins are often
associated with changes in the protein functions. PTM results
in changes in the molecular weight, charge, and hydrophobic-
ity of proteins. The modified subproteomes can be studied at
both protein and peptide levels using different gel-based anal-
ysis techniques, as well as MS. Glycosylation and phosphory-
lation are two common forms of modifications affecting about
40% and 60% of the seed proteins. Glycosylation events are
characteristic on all secreted and membrane proteins, while
phosphorylation is often linked to proteins related to enzyme
metabolism, stress responses, and growth. Storage proteins of
the cereal seeds are rich in amino acids like proline or gluta-
mine. Deamidation of glutamine-containing proteins into glu-
tamic acid results in higher water-binding capacity. Polymer
formation of storage proteins in wheat gluten may result from
PTM events, by which sulfur-containing amino acids are
152 PROTEINS | Proteomic Analysis Relevant to Cereal Grains
bound to each other via disulfide bonds, so that a stabile
chemical interaction is formed. These abilities of cereal and
especially wheat grain proteins are important in the structure
function relationship analysis of wheat gluten for food prod-
ucts and also for nonfood uses.
Computational and Structural Proteomics
Structural proteomics deals with the complex information hid-
den in the amino acid sequence of a protein. It determines how
the order and characteristics of the constituent amino acids
influence the secondary and tertiary structure of a protein
and how these features determine the ability of these proteins
to function as interacting partners in protein complexes. The
major challenge of structural proteomics is to assign structure
data determined by x-ray or NMR crystallography or in silico
modeled to protein sequences obtained from genome and
transcriptome sequencing projects by the use of high-
throughput characterization of the 3-D structure. Structural
proteomic studies of cereal grains can be used to analyze
enzyme complexes present in the seed, to identify receptor
functions by the analysis of different proteinligand interac-
tions, to predict how pathogens (including viruses, fungi, or
insects) interact with the host plant, or to identify and predict
allergen components present in the seed. Structural proteomics
strongly builds on computational biology utilizing the poten-
tial of sequence data analysis and 3-D modeling.
Computational proteomics is a term generally describing a
broad range of computational tools, including algorithms, bio-
informatics techniques, and data acquisition models to analyze
protein or peptide related sequence data. The collection and
evaluation of data from protein databases, combined with gel-
based and MS-based or immunoanalytical computational ana-
lyses, can deliver outputs in drug discovery, food-safety analyses,
and even the development of personalized medicine.
Improving on Current Proteome Technology
The successful completion of wheat genome and genome-
sequencing projects of related grain species will lead us to
better understand mechanisms involved in the stages of plant
development, such as seed development, maturation, seed
dormancy, and germination. Additional insights will come
from using genome sequences combined with genotype-level
information, plus additional achievements in protein separa-
tion and identification techniques. Results can be expected in
genotype-level complex information about what features are
responsible for specialized end-use quality attributes, for sus-
tainable quality under different environmental conditions, and
for the complexity of wheat-allergen proteins. To study these
mechanisms, projects need to involve multidisciplinary
approaches, such as genomics, proteomics, metabolomics,
and bioinformatics.
Currently, proteomics receives criticisms related to its low
reproducibility, evidenced by gel-to-gel variations between lab-
oratories. By the standardization of extraction and separation
protocols, these issues will be improved. Identification of
weakly expressed molecules via MS remains problematic,
although techniques such as protein and peptide chips
combined with Western blotting may be used to overcome
this challenge. The power of proteomics used for the identifi-
cation and characterization of protein components in the grain
is greatly magnified when combined with other postgenomic
techniques such as metabolomics. Such multidisciplinary ana-
lyses, including tools of proteomic, metabolomic, and bioin-
formatic analyses, can provide direct evidence of the effects of
protein expression.
The next major challenge toward this aim is to identify the
constituents of the cereal grain proteome in order to under-
stand the cereal genome. Of particular importance will be the
deciphering of protein alterations between normal and stressed
conditions to permit the identification and prioritization of
agronomically important targets. Proteome analysis aims to
identify changes not only in protein expression but also in
PTMs, proteinprotein interactions, cellular and subcellular
distribution, and temporal patterns of expression.
The power of proteomics as a tool for the discovery of
biological pathways and processes related environmental
changes is now well established. Indeed, proteomics has
already uncovered many potential targets for specialized qual-
ity purposes or different biotic and abiotic stress conditions.
The current era of proteomics is now beginning to investigate
how this technology can serve the breeders, the food industry,
and the consumer for high-throughput identification and
prognostic applications.
Exercises for Revision
Develop an extended definition of the proteomics concept,
including mention of the genome, growth conditions, tis-
sue identity, stage of maturity, and functional relationships.
Explain the relative advantages and disadvantages of gel-
based and gel-free technologies for proteome analysis.
Which is better suited for polypeptide identification and
for eventually elucidating the functions of the many
polypeptides?
Several distinct versions of proteomics are described:
expression, structural, functional, posttranslational
proteomics, etc. Explain the differences between these
concepts and their successive roles in understanding the
sequence of events from genes to mature grain.
Describe some practical applications of proteomics in grain
science.
Construct a diagram (and explanation) showing relation-
ships between genomics, various forms of proteomics, and
other -omics. Describe their relevance to grain science.
Consider thiolomics and zymomics, as examples (but not
economics). The following reference provides an introduc-
tion to zymomics: Kosar-Hashemi B, Irwin JA, Higgins J,
Rahman S, and Morell MM (2006) Isolation, identification,
and characterization of starch-interacting proteins by 2-D
affinity electrophoresis. Electrophoresis 27: 18321839.
Exercises for Readers to Explore the Topic Further
It is stated that the results of proteomic analysis of grain
proteins have wide-ranging implications for research on
 PROTEINS | Proteomic Analysis Relevant to Cereal Grains
structurefunction relationships for grain proteins. Find
out about some examples of proteomics being used to
demonstrate how specific proteins influence significant
aspects of grain development and utilization.
Develop some diagnostic applications of proteomic analy-
sis, based on the knowledge that specific polypeptides dis-
tinguish one cereal variety from others (variety
identification), or one grain tissue from another (milling)
or one grain genus from another, or grain tissue from that
of a predator.
Investigate applications of proteomics in plant breeding to
achieve aims such as improved pathogen resistance and
better grain quality.
Construct a diagram (and explanation) showing relation-
ships between genomics, various forms of proteomics, and
other -omics. Describe their relevance to grain science.
Consider thiolomics and zymomics, as examples (but not
economics). The following reference provides an introduc-
tion to zymomics: Kosar-Hashemi B, Irwin JA, Higgins J,
Rahman S, and Morell MM (2006) Isolation, identification,
and characterization of starch-interacting proteins by 2-D
affinity electrophoresis. Electrophoresis 27: 18321839.
See also: Breeding of Grains: Wheat Breeding: Exploiting and
Fixing Genetic Variation by Selection and Evaluation; Genetics of
Grains: Development of Genetically Modified Grains; Genomics of
Food Grains; Wheat Genetics and Genomics; Grain Marketing and
Grading: Identification of Varieties of Food Grains; Proteins: Protein
Synthesis and Deposition; The Protein Chemistry of Cereal Grains;
Thiolomics of the Gluten Protein Network of Wheat Dough; The
Basics: Grain and Plant Morphology of Cereals and how characters
can be used to identify varieties; Grain: Morphology of Internal
Structure; Wheat Processing: The Gluten Proteins of the Wheat
Grain in Relation to Flour Quality.
Further Reading
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Zivy M, and Damerval C (eds.) Plant Proteomics Methods and Protocols. Methods
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Camafeita E, Sols J, Alfonso P, Lopez JA, Sorell L, and Mendez E (1998) Selective
identification by matrix-assisted laser desorption/ionization time-of-flight mass
spectrometry of different types of gluten in foods made with cereal mixtures. Journal
of Chromatography A 823: 299306.
Colinge J and Bennett KL (2007) Introduction to computational proteomics. PLoS
Computational Biology 3(7): e114. http://dx.doi.org/10.1371/journal.
pcbi.0030114.
Cramer R and Westermeier R (eds.) (2012) Difference Gel Electrophoresis (DIGE).
Methods and Protocols. In: Methods in Molecular Biology, vol. 854. Totowa, NJ:
Humana Press.
Dupont FM, Vensel WH, Tanaka CK, Hurkman WJ, and Altenbach SB (2011)
Deciphering the complexities of the wheat flour proteome using quantitative two-
153
dimensional electrophoresis, three proteases and tandem mass spectrometry.
Proteome Science 9: 10. http://dx.doi.org/10.1186/1477-5956-9-10.
Finnie C, Sultan A, and Grasser KD (2011) From protein catalogues towards targeted
proteomics approaches in cereal grains. Phytochemistry 72: 11451153.
Gao L, Wang A, Li X, et al. (2009) Wheat quality related differential expressions of
albumins and globulins revealed by two-dimensional difference gel electrophoresis
(2-D DIGE). Journal of Proteomics 73: 279296.
Gao L, Ma W, Chen J, et al. (2010) Characterization and comparative analysis of wheat
high molecular weight glutenin subunits by SDS-PAGE, RP-HPLC, HPCE, and
MALDI-TOF-MS. Journal of Agricultural and Food Chemistry 58: 27772786.
Hashiguchi A, Ahsan N, and Komatsu S (2010) Proteomics application of crops in the
context of climatic changes. Food Research International 43: 18031813.
Jorrin-Novo JV, Komatsu S, Weckwerth W, and Wienkoop S (eds.) (2014) Plant
Proteomics Methods and Protocols. In: Methods in Molecular Biology, vol. 1072.
Totowa, NJ: Humana Press.
Juhasz J, Moolhuijzen P, Bellgard M, Appels R, and Bekes F (2013) Wheat grain
proteomics for the food industry. In: Toldra F and Nollet LML (eds.) Proteomics in
Foods. Principles and Applications. Food Microbiology and Food Safety,
2: pp. 341379. New York: Springer.
Kav NNV, Srivastava S, Yajima W, and Sharma N (2007) Application of proteomics to
investigate plant-microbe interactions. Current Proteomics 4: 2843.
Manjasetty BA, Bussow K, Panjikar S, and Turnbull AP (2012) Current methods in
structural proteomics and its applications in biological sciences. 3 Biotech 2(2):
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Martnez-Maqueda D, Hernandez-Ledesma B, Amigo L, Miralles B, and Gomez-Ruiz J
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digestion. In: Toldra F and Nollet LML (eds.) Proteomics in Foods. Principles and
Applications. Food Microbiology and Food Safety, vol. 2, pp. 2150. New York:
Springer.
McLaffety FW, Breuker K, Jin M, et al. (2007) Top-down Ms, a powerful complement to
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Miernyk JA and Hajduch B (2011) Seed proteomics. Journal of Proteomics
74: 389400.
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Skylas DJ, Mackintosh JA, Cordwell ST, et al. (2000) Proteome approach to the
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Tasleem-Tahir A, Nadaud I, Girousse C, Martre P, Marion D, and Branlard G (2011)
Proteomic analysis of peripheral layers during wheat grain development.
Proteomics 11: 371379.
Relevant Websites
http://expasy.org Proteomics tools.
http://propepper.net ProPepper is a protein, peptide and epitope analysis suitable for
cereal prolamin analysis.
www.uniprot.org Uniprot is a comprehensive database and analysis tool used protein
science.
http://wiley-vch.de Proteomics is a scientific journal specializing in articles on
proteomics.
 Thiolomics of the Gluten Protein Network of Wheat Dough
F Bonomi and S Iametti, University of Milan, Milan, Italy
2016 Elsevier Ltd. All rights reserved.
Abbreviations
2-ME 2-Mercaptoethanol
5-IAF 5-Iodoacetamidofluorescein
DTT Dithiothreitol
HMW-GS High-molecular-weight glutenins
Topic Highlights
Cysteine thiols and disulfides play key roles in the forma-
tion and stabilization of gluten network.
Disulfides may be inter- or intramolecular.
Intramolecular disulfides are typical of gliadins; inter-
molecular, of glutelins.
The pattern of free cysteines and of disulfides is typical of
individual components in both gliadins and glutelins.
Accessibility of cysteine thiols to suitable reagents depends
on structural features of the protein(s).
Denaturing agents cause changes in the accessibility of
cysteine thiols to specific reagents.
Accessible thiols in individual polypeptides may be studied
after they are covalently labeled with a fluorescent reagent.
Separation of covalently labeled polypeptides by 2-D elec-
trophoresis allows to identify species- and variety-specific
accessibility patterns.
Image analysis of thiol-stained and protein-stained 2-D
electrophoretic patterns forms the basis of thiolomics.
Thiolomics combines data on the occurrence/abundance of
individual proteins with those on the accessibility of their
cysteine thiols in the presence/absence of chaotropes.
Learning Objectives
To understand the role of thiols and disulfide in the estab-
lishment of intra- and intermolecular covalent bonds
among cereal proteins.
To acquire knowledge on how individual proteins in cereals
may contribute to process-induced formation of a protein
network through disulfide exchange reactions.
To get acquainted with methodologies used to study the
accessibility of free thiols in cereal proteins and process-
related changes in their accessibility pattern.
To get an overview of how proteomics-based approaches
can be combined with chemical labeling of accessible
thiols, in order to evaluate the possible contribution of
individual species to the desired product properties.
Relevance of Thiols and Disulfides to
ProteinProtein Interactions
Formation of a viscoelastic network involving wheat gliadins
and glutelins (and similar proteins in related cereals) is a
154 Encyclopedia of Food Grains, Second Edition
LMW-GS Low-molecular-weight glutenins
PAGE Polyacrylamide gel electrophoresis
SDS-PAGE Sodium dodecyl sulfate polyacrylamide
gel electrophoresis
crucial step in the production of an extremely diverse variety
of common wheat-based foods. Gliadins and glutelins repre-
sent two classes of water-insoluble proteins found in wheat
flour and semolina. The so-called gluten network formed by
these two protein classes contributes to these systems an array
of features that have a deep impact on the consumer appreci-
ation of the final product, be that due to retention of gas
bubbles in bread and baked products or to entrapment of
swollen starch in pasta.
The interactions leading to the formation of the viscoelastic
network in gluten are of two main types, namely, (1) hydro-
phobic contacts among proteins (or within individual
proteins) and (2) intra- and intermolecular disulfides. Most
often, these bonds are present in the original molecules, but
they are rearranged in processes that require protein structural
flexibility (made possible by the addition of water prior to the
mixing steps) and the action of shear forces that act as
mechanical denaturants during mixing. Covalent bonds
form as a consequence of oxidation of spatially closed cysteine
thiols to form a disulfide bridge or more commonly in the
so-called disulfide exchange reactions, which in turn require
the presence of free and accessible cysteine thiols and of a
preexisting disulfide in specific proteins. Disulfide exchange
among proteins upon structural modification is a common
occurrence in many processed foods, from hard-boiled eggs
to heat-treated milk.
Given the relevance of disulfides and thiols to gluten prop-
erties, chemical and biochemical oxidants and reductants have
often been used as ameliorants of wheat flour. Oxidants (such
as bromate/iodate or hydrogen peroxide that may also be
produced in situ by appropriate enzymes) promote the forma-
tion of disulfides, whereas reducing agents (such as ascorbate,
cysteine, and free or protein-bound glutathione) increase the
number of free thiols and could facilitate thioldisulfide
exchange reactions.
In common practice, the amount of proteins in wheat flour
or semolina and the glutelin/gliadin ratio in wheat flour are
considered key parameters to predict their best possible use.
Proteomics-based studies have provided a comprehensive and
detailed view of the protein patterns in wheat, but attempts at
finding correlations with the processing performance seldom
indicated evident connections. Also, environmental and agro-
nomic issues may affect either protein expression or protein
structure after proteins are deposited in the seed.
Among the many other parameters that need to be consid-
ered in a comprehensive approach to protein quality are those
http://dx.doi.org/10.1016/B978-0-12-394437-5.00108-X
 PROTEINS | Thiolomics of the Gluten Protein Network of Wheat Dough
related to structural issues in these proteins. These include (1)
the role of noncovalent (mostly hydrophobic) interactions
in network formation; (2) the availability of thiols and disul-
fide to exchange reactions, as determined by overall and
local protein flexibility, in turn related itself to protein solv-
ability and to the intensity of physical deformation; and
(3) the possible role of low-molecular-weight thiols and
of low-abundance thiol-rich proteins in facilitating thiol
disulfide exchange reactions.
Structural studies on proteins that cannot be solubilized
without resorting to the use of denaturants (alcohols or deter-
gents) are intrinsically complicated. Methods capable of eval-
uating protein surface hydrophobicity in systems made up of
water-insoluble proteins have been developed and applied to
protein characterization in cereal-based starting materials and
products without resorting to protein separation that necessarily
uses solvent systems that affect protein structure and interactions
or to standard proteomics or peptidomics approaches, in properties are reportedly strongly influenced by their relative
which a combination of separative techniques is used to clarify
the whole pattern of proteins and peptides and to quantitate
their relative abundance. Protein solubility in the absence/pres-
ence of chaotropes and in the absence/presence of disulfide
reductants, such as dithiothreitol (DTT) and 2-mercaptoethanol
(2-ME, also known as beta-mercaptoethanol), has been used to
study hydrophobic and covalent interactions among proteins
and their impact on the rheological properties of dough. These
properties are reportedly affected by agents that block the
reactivity of thiols in wheat proteins, thus preventing many of
the permanent structural changes that occur during further
processing.
The thiolomics approach described in what follows aims
at combining information about the amount of thiols present
(and accessible under native or denaturing conditions) in indi-
vidual protein components in a complex system such as grain
proteins. This approach offers the advantage of combining
information about thiols on individual proteins along with
their location on specific components (which may be identi-
fied by further proteomics analysis), thus contributing to a
more comprehensive assessment of the molecular determi-
nants of grain performance.
Protein Thiols in Cereal Systems
Wheat proteins will form the basis of the discussion here, as
the occurrence and relevance of thiols in small molecules (the
so-called low-MW thiols that include free and protein-bound
glutathione and free cysteine itself) and in the water-soluble
albumin/globulin components of flours will be discussed
elsewhere. It has to be noted that some of the processes used
to convert some cereals into foods (including fermentation,
as in the case of sorghum) result in a significant decrease of
the total content of reactive cysteines that appear to be pref-
erentially metabolized by the involved microorganisms.
These effects on protein thiols are not as evident in other
pretreatments of cereal and noncereal grains (e.g., germina-
tion and/or malting).
As summarized in Figure 1, wheat gliadins are monomeric
proteins with a relevant role in the development of the gluten
intermolecular network through the formation of hydrogen
bonds, of hydrophobic bonds between nonpolar amino acid
155
side chains, and of thioldisulfide exchange reactions.
Gliadins also interact with the flour lipids or with added
lipids, and there is a general consensus on their role in deter-
mining dough extensibility. Gliadins have molecular masses
ranging from 30 to 45 kDa and had been subdivided into a/b,
g, and o fractions, according to their mobility in polyacryl-
amide gel electrophoresis (PAGE) at acid pH. Alpha- and
b-gliadins are currently considered a single gliadin (a/b
type) and contain six Cys residues in their C-terminal
domain. Eight Cys residues are found in the C-terminal
domain of g-gliadins, whereas no cysteine is present in
o-gliadins.
The glutenin fraction in wheat consists of protein aggregates
from about 60 000 to more than 10 millions Da. Glutenin sub-
units have been divided into high-molecular-weight glutenins
(HMW-GS) and low-molecular-weight glutenins (LMW-GS).
HMW-GS represent 510% of gluten proteins, but dough
abundance. The bread-making quality of wheat genotypes
appears to be determined by the expression of specific HMW-
GS genes. The N- and C-terminal domains in HMW-GS contain
cysteine residues that form inter- and intrachain disulfide
bonds, whereas the central domain comprises repeated
sequences that may promote intermolecular hydrogen bonding.
In particular, the x-type subunit includes four cysteines; three at
the N-terminal and one at the C-terminal. The y-type subunit
has seven cysteines; five in the N-terminal domain, one within
the repetitive domain, and one close to the C-terminus. These
proteins promote the formation of larger glutenin polymers by
forming intermolecular disulfides with other glutenins.
LMW-GS are classified into B (4251 kDa), C (3040 kDa),
and D types (5570 kDa), the B type being the most abundant.
D-type LMW-GS correspond to a o-gliadin-like protein with an
additional cysteine that allows interchain disulfide bond for-
mation in the glutenin polymer. C-type LMW-GS have a struc-
ture similar to a/b- and g-gliadin but contain an extra unpaired
cysteine residue thought to be involved in an interchain disul-
fide. The presence of an odd number of cysteines makes the
C- and D-type LMW-GS behave as terminators of the growing
glutenin polymer chains, with a presumed negative effect on
flour quality.
B-type LMW-GS are subdivided in LMW-s, LMW-m, and
LMW-i types, on the basis of the first amino acid in the mature
polypeptide being Ser, Met, or Ile, LMW-s-type subunits being
the most abundant. The sequences of LMW-s and LMW-m
consist of a short N-terminal region including the first cysteine,
a glutamine-rich repetitive domain, and a C-terminal region
containing seven cysteines. Mature LMW-m and LMW-s con-
tain eight cysteines, of which the first and seventh are involved
in intermolecular disulfide bonds, whereas the others partici-
pate in intramolecular bonds. LMW-i contain the typical
eight cysteine residues, all in the C-terminal domain and all
involved in interchain disulfide bonds. The ability of LMW to
form a variable number of intermolecular disulfide bonds
defines their role as chain extenders or chain terminators.
B-type LMW-GS allow the extension of glutenin polymers
(chain extenders). LMW-GS of the C type and D type behave
as chain terminators, in what they block further extension
of the disulfide network, as they lack the additional free
cysteine residues required for formation of further interchain
linkages.
156 PROTEINS | Thiolomics of the Gluten Protein Network of Wheat Dough

N-TER REP C-TER /

2 3 4 6
C C C C

C1 C5

N-TER REP C-TER

C2 C3 C4 C5 C6 C8

C1 C7

Gliadins
HMW
N-TER REP C-TER x-type
1 2 3
C C C C4

N-TER REP C-TER Dx5

1 2 3 4
C C C C C5

N-TER REP C-TER y-type

1 2 3 4 5 6 7
CC C C C C C

SH SH

LMW C 1
C7

N-TER REP C-TER s-and m-type

3 4 6 8
C C C C
C2 C5

N-TER REP C-TER i-type

C2 C3 C4 C5 C6 C8
C C

SH SH

Glutenins
Figure 1 Location of cysteine residues and of disulfide bonds in wheat gliadins and glutelins.

Thiol Accessibility as a Protein Structural Probe colored thionitrobenzoate anion upon reacting with an SH
group. This method can be applied in principle regardless
Accessibility of thiols to common colorimetric or fluorescent of the solubility of the proteins being investigated, but it is not
reagents has been studied as a function of added denaturing suitable for permanent (i.e., covalent) labeling of thiols, in
agents (urea, thiourea, guanidinium ions, and detergents) and view of the lability of the resulting mixed disulfide.
of reaction time to assess overall and local rigidity of proteins Other reagents can form much more stable bonds with
in either the starting material or the finished products. This protein thiols, generating species that may resist the chemically
approach could to some extent be combined with solubility severe conditions used for resolving individual components in
and compositional data, in an attempt to gather a unifying
vision of individual molecular features that contribute to the O OH
O
properties of these systems.
The approach presented here relies on the observation that N+
O O
thiols buried within the structure of a protein (or a protein S
aggregate) often become available to suitable reagents only HO S
O
upon protein denaturation by physical or chemical agents. +
N
Accessibility of the target thiol(s) is thus determined by steric
O
factors, in particular when bulky reagents are used. From a
methodological standpoint, readily accessible thiols may be O
NO2 NO2 NO2
SH + S OH S +
assessed in the absence of suitable protein denaturants, R HO S
R S
OH
HS
OH
O2N O O O
whereas the addition of denaturing agents should permit the
detection of all free thiols in the investigated proteins. Figure 2 Structure of 5,50 -dithiobis-(2-nitrobenzoic acid) (DTNB)
Simple spectrophotometric assays rely on using 5,50 -dithio- (Ellmans reagent) and its reaction with thiols, leading to the formation of
bis-(2-nitrobenzoic acid) (DTNB, see Figure 2) that releases a the yellow-colored (A412 13 700) thionitrobenzoate anion.
 PROTEINS | Thiolomics of the Gluten Protein Network of Wheat Dough 157

protein mixtures. There are three fundamental types of chemical
reactions that can be used for this purpose (see Figure 2),
involving (1) mercuric derivatives, (2) N-ethylmaleimide
derivatives, (3) iodinated compounds, such as monoiodobimane
or iodoacetamide derivatives. Selected examples of compounds
in each of these classes of reactivity are presented in Figure 3.
Mapping the Pattern of Thiol Exposure in Individual
Cereal Proteins (Thiolomics)
The possibility of covalently labeling any exposed thiol prior to
separation of individual polypeptides has been exploited to
assess the pattern of thiol exposure also in the insoluble com-
ponents of wheat flours and semolina. By introducing dena-
turing and disulfide-reducing steps at appropriate points in the
thiol-labeling procedure, it is possible to use 2-D electropho-
resis techniques to obtain a visually informative thiolomics
map of the conditional accessibility of thiols in individual
proteins in the materials under investigation. These maps pro-
vide information as for the type of proteins where thiols are
available and on the relative abundance of proteins and reac-
tive thiols on them (Figure 4).
In the original procedure, thiols in wheat flour or semolina
proteins were covalently labeled by 5-iodoacetamid-
ofluorescein (IAF) in the absence or in the presence of 4 M
urea, allowing to evaluate thiols accessibility before and after
protein unfolding and as related to the dissociation of non-
covalently linked protein complexes. Proteins labeled under
either condition were then separated by SDS-PAGE and/or by
2-D electrophoresis. Visual inspection and quantitative com-
parison of the maps for proteins and for accessible thiols in
different and variously treated starting materials were then
carried out to identify specific proteins or protein classes
whose thiol functions may specifically contribute to specific
traits of the material.
Only readily accessible protein thiols should be labeled in
flour samples treated with IAF in the absence of urea, whereas
O that covalent binding of the probe may introduce some alter-
Iodoacetamide
I ations in the pI of labeled proteins, although the impact of this
NH2
O O
H3C
N
CH3 Monobromobimane
N analyzed by standard image analysis techniques that may be
H3C CH2Br
O ground) and to transform halftone images into very informa-
N-Ethylmaleimide
N
O tive basis) compositional and/or structural differences among
O CH3
Hg
Phenylmercury acetate
O quently be stained for proteins by any of the procedures used
Figure 3 Examples of common thiol reagents suitable for covalent
labeling of proteins.
additional thiols buried in the hydrophobic regions of proteins
or at hydrophobic interfaces among proteins should become
available to the labeling agent if denaturants were added prior
to IAF. As stated earlier, measurements based on thiol accessi-
bility may be carried out regardless of whether the involved
proteins are soluble or insoluble (as is the case for most of
wheat proteins) in the conditions used for labeling.
As indicated by electrophoretic separation of IAF-labeled
wheat proteins in the presence of sodium dodecyl sulfate
(SDS-PAGE; see an example in Figure 5), treatment with IAF
in the presence of urea gave the expected increase in the inten-
sity and number of fluorescent bands, as a consequence of
increased thiol exposure in the presence of the chaotrope.
The SDS-PAGE tracings in Figure 5 also allow to appreciate
some specificity of the labeling procedure in wheat flour and in
semolina and a different sensitivity of otherwise similar pro-
teins in each of the samples to the presence of urea during the
labeling step. In the case of soft wheat, labeling in the presence
of urea apparently modifies only the intensity of the fluores-
cence associated with all protein bands, and no additional
bands were present in the sample labeled in the presence of
urea. In the case of durum wheat semolina, the fluorescence
intensity of specific protein bands was sensibly increased when
labeling was carried out in the presence of urea, and the effect
of the denaturant on the accessibility of thiols was most evi-
dent in the largest polypeptides. This may be taken as an
indication of the presence of multiple thiols on the involved
proteins and of changes in their accessibility to the reagent in
the presence/absence of urea.
Given the possible ambiguities associated with identifica-
tion of specific components from SDS-PAGE gels, 2-D electro-
phoretic maps were also obtained, similar to those used in
classic proteomics studies. In this technique, proteins are
made soluble by unfolding in urea and reduction with DTT
and are separated first according to their isoelectric points (pI)
in an immobilized pH gradient. The second run is performed
in an orthogonal direction with respect to the first separation
step and separates them by molecular mass. A comparison of
protein-stained 2-DE maps obtained in the presence and
absence of treatment with the fluorescent thiol probe confirms
substitution on the acidic proteins that are predominant in
wheat does not impair analysis of the resulting maps.
Typical 2-D maps of the IAF-labeled polypeptides in various
wheat samples are shown in Figure 6. These maps can be easily
used to convert positive and negative images (the original
fluorescence maps had bright spots against a black back-
tive and/or easily read landscape plots (see the example
presented in Figure 7). This type of graphical rendition
makes it easier to appreciate (both visually and on a quantita-
samples as reflected in their thiol accessibility maps.
It has to be noted that the very same 2-D gel can subse-
in standard proteomics studies (see Figure 8). Image analysis
of the protein-stained gel can provide additional information
on not only the occurrence and abundance of specific protein
158 PROTEINS | Thiolomics of the Gluten Protein Network of Wheat Dough

O OH
O
O
I

N O
H

OH

Iodoacetamide
fluorescein (IAF)

Cysteine
thiols
Interprotein
disulfide bonds

Hydrophobic
contact regions

Native, no Covalently bound Denaturant

denaturant fluorophor present

Labeled species present after denaturing/reducing treatments

Figure 4 A graphical rendition of the events related to covalent binding of the fluorescent probe IAF to wheat protein thiols in the absence/presence
of denaturants.

Urea Urea Urea Urea No urea + urea

+ + + +

Figure 6 2-DE pattern of IAF-labeled proteins in soft wheat flour.

Soft wheat Semolina
Figure 5 SDS-PAGE patterns for proteins reacted with IAF under
various conditions. In each figure, the left panel is Coomassie blue-
stained, and the right panel is the corresponding fluorescence image.
Letters on individual lanes indicate whether the treatment with IAF was
carried out in the presence () or in the absence () of 4 M urea.
components but also comparison with the fluorescence label-
ing maps allowing information to be gathered on the presence
and exposure of thiol groups in each of them. Furthermore,
since labeling with IAF does not to compromise identification
of specific protein components by standard mass spectrometric
Labeling with IAF was carried out in the absence or presence of urea, as
indicated. Images are digitally generated negatives from the original
fluorescence maps. Dashed areas indicate artifacts.
techniques, it is possible to identify individual proteins
present with exposed or buried thiols in materials of different
technological performances or in products obtained through
different processes.
In conclusion, thiolomics represents along with other
omics approaches a potentially powerful tool to assess
features of grain proteins that may be relevant to their techno-
logical performance. In comparison with approaches based on
the analysis of individual proteins, it offers the advantage of
 PROTEINS | Thiolomics of the Gluten Protein Network of Wheat Dough
(a) Hard wheat
Figure 7 Landscape plots obtained after image analysis of IAF-labeled 2-D electrophoretic patterns.
SilverCoomassie Fluorescence
Figure 8 2-DE pattern of IAF-labeled proteins in hard wheat flour.
Treatment with IAF was carried out in the presence of urea. The 2-DE gel
was stained for proteins with silverCoomassie blue (left) after
acquiring the fluorescent image (right). Green arrows indicate examples
of proteins that were present in marginal amounts but had a high
content of reactive thiols. Red arrows indicate examples of proteins
present in substantial amounts, but with no reactive thiols.
offering a comprehensive view of both the protein and the
thiol pattern in a complex system and of discriminating
between protein structures of different accessibilities.
Exercises for Revision
Define the relevance of covalent and noncovalent interac-
tions among proteins as for establishing a stable network.
Why is the pattern of exposed thiols expected to be different
in native and denatured proteins?
Do you expect protein thiols to be more accessible in pizza
or in spaghetti?
What could happen if all the free thiol groups in wheat
proteins were blocked (e.g., by chemical reagents) prior to
mixing cereal flours?
Provide a rationale for the observation that the pattern of
thiol-labeled proteins in a 2-DE analysis of wheat flour does
not overlap the pattern of proteins.
Why is the pattern of thiol-labeled proteins different when
the necessary dissociating step is carried out prior to the
labeling procedure?
Exercises for Readers to Explore the Topic Further
Extension of thiol accessibility/labeling studies to systems
containing proteins other than those of cereal origin may
159
(b) Soft wheat
provide interesting insights into their participation in the
establishment of an interprotein network. Discuss.
Thiol accessibility/labeling studies may also help to under-
stand the behavior of those systems (e.g., germinated grains
or fermented flours) where cereal proteins may undergo
proteolytic events and where a mechanical denaturation
step (as in mixing/kneading) is not present. Propose a
specific study to investigate this.
The thiolomics approach as reported here could be further
improved and adapted to studies on the relevance of
individual steps in specific processes (e.g., by using other
protein dissociating agents and/or other thiol-specific
reagents). Design an appropriate study.
See also: Proteins: Protein Synthesis and Deposition; Proteomic
Analysis Relevant to Cereal Grains; The Protein Chemistry of Cereal
Grains; Wheat Processing: Analysis of Dough Rheology in
Breadmaking; The Gluten Proteins of the Wheat Grain in Relation to
Flour Quality.
Further Reading
Belton PS (1999) On the elasticity of wheat gluten. Journal of Cereal Science
29: 103107.
Bonomi F, Mora G, Pagani MA, and Iametti S (2004) Probing structural features of
water-insoluble proteins by front-face fluorescence. Analytical Biochemistry
329: 104111.
Bonomi F, DEgidio MG, Iametti S, et al. (2012) Structurequality relationship in
commercial pasta: A molecular glimpse. Food Chemistry 135: 348355.
Bonomi F, Iametti S, Mamone G, and Ferranti P (2013) The performing protein: Beyond
wheat proteomics? Cereal Chemistry 90: 359366.
Field JM, Shewry PR, and Miflin BJ (1983) Solubilization and characterization of wheat
gluten proteins; correlations between the amount of aggregated proteins and baking
quality. Journal of the Science of Food and Agriculture 34: 370377.
Goesaert H, Brijs K, Veraverbeke WS, Courtin CM, Gebruers K, and Delcour JA (2005)
Wheat flour constituents: How they impact bread quality, and how to impact their
functionality. Trends in Food Science and Technology 16: 1230.
Iametti S, Marengo M, Miriani M, Pagani MA, Marti A, and Bonomi F (2013) Integrating
the information from proteomic approaches: A thiolomics approach to assess
the role of thiols in protein-based networks. Food Research International
54: 980987.
Joye IJ, Lagrain B, and Delcour JA (2009) Endogenous redox agents and enzymes that
affect protein network formation during breadmaking. Journal of Cereal Science
50: 110.
Mamone G, Addeo F, Chianese L, et al. (2005) Characterization of wheat gliadin
proteins by combined two-dimensional gel electrophoresis and tandem mass
spectrometry. Proteomics 5: 28592865.
160 PROTEINS | Thiolomics of the Gluten Protein Network of Wheat Dough
Mamone G, De Caro S, Di Luccia A, Addeo F, and Ferranti P (2009) Proteomic-based
analytical approach for the characterization of glutenin subunits in durum wheat.
Journal of Mass Spectrometry 44: 17091723.
Morel MH, Redl A, and Guilbert S (2002) Mechanism of heat and shear mediated
aggregation of wheat gluten upon mixing. Biomacromolecules 3: 488497.
Shewry PR, Tatham AS, Forde J, Kreis M, and Niflin BJ (1986) The classification and
nomenclature of wheat gluten protein: A reassessment. Journal of Cereal Science
4: 97106.
Shewry PR, Halford NG, Tatham AS, Popineau Y, Lafiandra D, and Belton PS (2003) The
high molecular weight subunits of wheat glutenin and their role in determining
wheat processing properties. Advances in Food and Nutrition Research
45: 219302.
Singh H and MacRitchie F (2001) Application of polymer science to properties of
gluten. Journal of Cereal Science 33: 231243.
Skylas DJ, Van Dyk D, and Wrigley CW (2005) Proteomics of wheat grain. Journal of
Cereal Science 41: 165179.
Veraverbeke WS and Delcour JA (2002) Wheat protein composition and properties of
wheat glutenin in relation to bread-making functionality. Critical Reviews in Food
Science and Nutrition 42: 179208.
Wieser H (2007) Chemistry of gluten proteins. Food Microbiology 24: 115119.
 Carbohydrates
Contents
Carbohydrate Metabolism
Starch: Chemistry
Starch: Synthesis
Starch: Starch Architecture and Structure
Starch: Analysis of Quality
Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
Grains other than Cereals: Non-starch Polysaccharides
Resistant Starch and Health
Health Effects of b-Glucans Found in Cereals
Glycemic Index, Glycemic Load, and Their Health Benefits
Carbohydrate Metabolism
RN Chibbar, S Jaiswal, M Gangola, and M Baga, University of Saskatchewan, Saskatoon, SK, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Carbohydrates are one of the major components of food
grains.
Carbohydrates are mainly categorized on the basis of their
degree of polymerization, structure, and function.
Biosynthesis of different sugars occurs in different plant cell
organelles.
Carbohydrates support seed germination and early seedling
development.
Carbohydrates, besides being an energy source, also pro-
vide structural integrity to plant cells.
Carbohydrate biosynthesis can be targeted to improve food
grain quality for human consumption.
Carbohydrates are also involved in abiotic stress tolerance.
Learning Objectives
To achieve understanding of the structural diversity of grain
carbohydrates, their biosynthesis, storage in grains, and
mobilization during seed germination.
To appreciate the influence of structural diversity of grains
on their end use for human food, animal feed, and indus-
trial applications.
Introduction
Carbohydrates are one of the most important nutrients for
both plants and animals. In plants, carbohydrates constitute
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00089-9
two-thirds to three-quarters of dry matter and are the major
components of storage organs. Cellulose, the most abundant
plant carbohydrate, provides structural integrity to plants and is
used for fiber and fuel by humans. In seeds, carbohydrates
provide energy and substrates required for the germination
and initial growth of seedlings. Carbohydrate in the form of
starch is the major reserve polysaccharide present in grains.
Carbohydrates like starch and cellulose form the basis of several
important industries in the food, feed, and fuel sectors. They
also provide renewable and environmentally friendly raw mate-
rials for industrial applications such as biodegradable plastics,
adhesives, and ethanol-based fuels. With the development
of genetic modification technologies, the biosynthesis of car-
bohydrates can be altered in planta to generate novel products
for various food and industrial applications. This article will
describe the structure, occurrence, and distribution of some of
the common storage and structural carbohydrates present in
the grains of major crops. The biochemical reactions occurring
during the biosynthesis of carbohydrates and their utilization
during seed germination will be discussed.
Carbohydrates
Definition and Classification
The term carbohydrate, or hydrates of carbon, is derived from
its basic elemental formula [Cx(H2O)y], in which carbon is
joined to hydrogen and oxygen present in the same ratio as in
water. Chemically, carbohydrates are polyhydroxy aldehydes or
ketones, their simple derivatives, or their polymers. Carbohy-
drates in grains are classified based on their chemical structures
161
162 CARBOHYDRATES | Carbohydrate Metabolism

or their digestibility when consumed by humans as food or by a disaccharide. Upon hydrolysis of disaccharides, the glycosidic
livestock as feed. Based on the polymeric nature of carbohydrates, bond is split into yield the component monosaccharides. Sucrose
they are classified as monosaccharides, disaccharides, oligosac- and maltose are the most commonly occurring disaccharides
charides, and polysaccharides (Figure 1). Monosaccharides are in the majority of grains. Oligosaccharides include the sugars
the simplest of all sugars, contain at least three carbon atoms, and that contain 320 sugar molecules joined by glycosidic bonds.
are the building blocks of all carbohydrates. The most common The most common oligosaccharides are soluble a-galactosides,
monosaccharides in grains are the hexoses (fructose, glucose, and characterized by galactose moieties joined by a(1,6) linkages.
galactose) and the pentoses (arabinose and xylose). Two sugar In legumes, raffinose, stachyose, and verbascose are the most
moieties joined by a glycosidic bond (an oxygen bridge) form common oligosaccharides and constitute 618% of the legume

Monosaccharides

H H H
O H O O O
H H H
O O
O O O O O O O O
H H H H
O
H O
H H O O
H H
O O O O O O

O O H H
O O
H H H
H
-D-Glucose -D-Fructose -D-Xylose -D-Galactose -L-Arabinose

Disaccharides

H H
O H H
O O O
H H H
H H O O O O
O H
O O O
H H
O
O H O H
O O O O O
O O O O
O O O O O
H H H H H H
O H O O

O O O O O O O
H H H H H H H
O O O O O
H H H H H

Sucrose -D-Maltose Galactinol Lactose

Oligosaccharides
HO
HO H
H O O HO
OH OH O
H
H
H H O OH H H H H
O O
O H O O O O O
O O H HO O H
O O O OH
H O H O OH HO
O O O O O H O H O H
H O O H O
O O
HO O
O HO O O
OH
H O O H OH
O O O O HO O O
HO O
OH H O O
O O O O
H O O H O O
O
H O H H
H H H O O HO
O
H
Raffinose Manninotriose Stachyose H
HO
H Verbascose

HO H
O
H H H H
H O O O O O O
H H H H
O O O
O
O O O H O
O O O O O H
O H O
H O HO H O O
H
O O O H
O H H H
O O O HO O O
H OH O
H O H O
O O
H O O H
O O
O OH H
H O
O O
H O O
O O O H O
H O
O H OH H H
O
O O
O O
O H H
Lychnose H
O H Stellariose Mediose
H

Figure 1
 CARBOHYDRATES | Carbohydrate Metabolism 163

Polysaccharides
H
H
O
Main chain O OH
CH2OH CH2
CH2OH CH2
O O H O OH H O
O
OH O OH O O O O O
O O
OH OH H O OH
O OH CH2 CH2
-D-Glc(14) linkage n
H O O
H O-H H
HO O H
Amylose O O
O
O O
O
HO O H
CH2 CH2

O O

CH2OH CH2OH H H
O O
Branch
O OH O OH Cellulose units linked with inter and intra chain H bonds
OH OH O -D-Glc(16) branch point
CH2 CH2OH
CH2OH CH2OH
O O O O O OH
O O O
O
OH OH O O
O OH OH O O
O
OH OH OH
OH OH OH OH
OH OH OH
OH
Main chain
-D-Glc(14) linkage OH OH -1, 3-Xylan
n
Amylopectin

Fructans
CH2OH
O
OH
CH2OH
O
CH2OH CH2OH O OH
CH2OH CH2OH O CH2
O O
O OH
OH OH OH OH
CH2O CH2OH
O CH2O OH CH2OH
OH OH O OH (n)
OH CH2OH
OH (n) CH2 O
O OH
OH
CH2OH
O
OH OH
OH
Fructan-levan type
Fructan-inulin type

Sugar alcohols

O H
H
O O
H

O O
O O
H H
H H H H
H O O O O
O
O O
O O H H H
H H O O

O O O O O
O O O H H H H
H H
O
H O
H

Ciceritol Sorbitol Mannitol

Figure 1 (Continued) Classification and chemical structures of representative carbohydrates found in grains of crops. Structure database source:
http://www.ncbi.nlm.nih.gov/pccompound. CID number reference: a-D-glucose, 79025; b-D-fructose, 439709; a-D-xylose, 6027; a-D-galactose, 439357;
a-L-arabinose, 641630; sucrose, 5988; a-D-maltose, 439341; galactinol, 11727586; lactose, 6134; raffinose, 439242; manninotriose, 440660;
stachyose, 439531; verbascose, 441434; lychnose, 25079986; stellariose, 46926270; mediose, 46926113; ciceritol, 44237401; sorbitol, 5780; and
mannitol, 6251.

seed dry weight. The presence of these flatulence-causing Some of the most common polysaccharides present in grains
a-galactosides is one of the major reasons restricting consumption are starch, cellulose, and xylans such as pentosans, b-glucans,
of legume-based food products. The absence of a-galactosidase and arabinoxylans. In relation to human and animal nutrition,
in the intestinal tract of humans causes fermentation of carbohydrates are often classified as available and unavailable.
a-galactosides by intestinal bacteria that contribute to the flatu- The available carbohydrates represented by sugars and starches
lence effect. Polysaccharides are polymers with more than 20 are broken down and absorbed in the human and animal diges-
monosaccharides joined together by glycosidic linkages. These tive tracts to provide energy for the body. Structural cell wall
are complex molecules because of the diversity in monosaccha- carbohydrates such as cellulose, hemicellulose, and nonstarch
ride units and type of linkages present in the polysaccharides. complex carbohydrates, for example, xylans, are categorized as
164 CARBOHYDRATES | Carbohydrate Metabolism

Table 1 Chemical composition of seeds of some important cereal grains (Table 3). Chickpeas especially have a high concen-
grain crops tration of soluble sugars, of which sucrose is the major compo-
nent. Lentils and peas have higher concentrations of stachyose
Composition (percentage dry
and verbascose as compared with other legumes. Starch is the
weight)
major storage carbohydrate in cereal grains, where it forms two-
Crop Carbohydrate Protein Lipid thirds to three-quarters of the dry weight. However, in oats, the
starch concentration is slightly less, ranging from 50% to about
Cereal crops 60%. Legume seeds also contain 50% starch, except lupins
Wheat (Triticum aestivum) 82 14 2 and soybeans in which only 0.51.4% of the dry weight is starch.
Barley (Hordeum vulgare) 80 9 1 The structural carbohydrates or functionally unavailable carbo-
Corn (Zea mays) 84 10 5
hydrates (dietary fibers) range in distribution from slightly above
Rice (Oryza sativa) 88 8 2
Oat (Avena sativa) 67 28 1
5% in rice to almost one-third of the grain dry weight in barley.
Sorghum (Sorghum vulgare) 82 12 4 In grain legumes, chickpeas have a higher concentration of total
Rye (Secale cereale) 82 14 2 nonstarch polysaccharides, whereas the concentration of total
Teff (Eragrostis tef) 83 11 2.5 dietary fiber corresponds to that of other legumes (Table 3). The
Pearl millet (Pennisetum glaucum) 81 12.5 5 major nonstarch complex carbohydrate in grain legumes is cel-
Foxtail millet (Setaria italica) 82 9.9 2.5 lulose, which represents about one-eighth of the grain dry weight
Finger millet (Eleusine coracana) 88 7 1.7 in most of the grain legumes (Table 3).
Kodo millet (Paspalum 84 11.5 1.3
scrobiculatum)
Proso millet (Panicum miliaceum) 82 10.6 4 Metabolism
Canary (Phalaris minor) 61 20.3 11
Triticale (xTriticosecale) 72 13 2.1
Autotrophic plants have the ability to convert simple nutrients
Cereal like non-Poaceae family grains such as carbon dioxide, water, and inorganic ions into interme-
Buck wheat (Fagopyrum 83 11 2 diates leading to the biosynthesis of complex molecules, such as
esculentum) nucleic acids, proteins, lipids, polysaccharides, and secondary
Quinoa (Chenopodium quinoa) 78 14.5 5.2 metabolites needed for growth and development (Figure 2).
Pulse crops Through the process of photosynthesis, plant cells capture light
Lentil (Lens culinaris) 65 27 1 energy to fix reduced carbon dioxide and water into a simple
Pea (Pisum sativum) 68 27 2 carbohydrate backbone, CH2O. The reaction takes place in chlo-
Chickpea (Cicer arietinum) 65 23 5 roplasts, wherein CO2 entering the Calvin cycle is incorporated
Kidney bean (Phaseolus vulgaris) 70 24 2
into 3-phosphoglycerate (3-PGA) molecules, which are subse-
Mung bean (Vigna radiata) 69 26 1
Pigeon pea (Cajanus cajan) 65 21 1
quently converted to triose phosphates. In addition, the
Soybean (Glycine max) 32 38 20 absorbed light energy is used for extracting reducing equivalents
Cowpea (Vigna unguiculata) 60 24 1 from water and ATP production required for processes involved
Lupin (Lupinus albus) 37 38 20 in plant growth and development. The reactions associated with
Oilseed crops light absorption are diurnal, which results in large variations in
Rape (Brassica napus) 25 23 48 nutrients available to plants during light and dark periods. How-
Sunflower (Helianthus annuus) 48 20 29 ever, plants have developed considerable flexibility in their
Flax (Linum usitatissimum) 32 26 38 metabolism to cope with fluctuations in nutrient concentration.
Peanut (Arachis hypogaea) 25 27 45 Plant cells can be envisioned to have a series of metabolic inter-
Sesame (Sesamum indicum) 19 20 54
mediate pools, which are interconnected to each other through
Hemp (Cannabis sativa) 27 29 41
enzyme reactions or transport mechanisms that are often revers-
ible (Figure 2). According to the physiological need(s) of a plant,
unavailable carbohydrates because they are not assimilated by the metabolites can be added to or withdrawn from these pools. Two
digestive tract. The unavailable carbohydrates are also often metabolite pools of the reductive pentose phosphate pathway
referred to as dietary fibers. and its intermediates are (1) the triose phosphate (3-PGA and
dihydroxyacetone phosphate (DHAP)) and (2) the hexose phos-
phate (glucose 1-P, glucose 6-P, fructose 1-P, and ADP-glucose)
Distribution in Important Grains
pools (Figure 2).
In general, carbohydrates dominate as the major constituent of The triose phosphates are either transported by triose phos-
the dry weight of seeds (Table 1). However, in oilseed grains, phate transporters to the cytosol or converted to other phos-
such as Brassica spp., lipids and proteins exceed carbohydrates as phorylated compounds, such as fructose 6-phosphate, in the
the major storage components by comprising three-quarters of plastid. The plastidial fructose 6-phosphate is used for both
grain dry weight. In cereal grains, the total concentration of regenerating ribulose-1,5-bisphosphate and producing glucose
mono-, di-, and oligosaccharides varies from < 1% to 3% 1-phosphate. Glucose 1-phosphate is converted to ADP-
(Table 2). Corn kernels contain the highest percentage of glucose glucose by ADP-glucose pyrophosphorylase (AGPase), the
and fructose, whereas in barley, sucrose is the major free sugar. immediate precursor for starch biosynthesis. Starch biosynthe-
Wheat grains have the highest concentration of raffinose among sis occurs inside plastids, but its functionality depends on the
the cereal grains (Table 2). Legume crops in general have a higher plastid type (chloroplast/amyloplast) and source plant tissue.
concentration of di- and oligosaccharides as compared with Starch synthesized in the chloroplast during the day is known as
 CARBOHYDRATES | Carbohydrate Metabolism 165

Table 2 Carbohydrate composition of some cereal grains

Free sugar (%) Starch (%) Nonstarch complex carbohydrates (%)

Total
dietary
Crop Total Glucose Fructose Sucrose Raffinose Total Amylose fiber Pentosan b-Glucan Cellulose

Wheat 1.41.3 0.020.03 0.020.04 0.570.80 0.540.70 6372 2328 14.6 6.6 1.4 2.0
Barley 2.03.0 0.10.2 0.1 1.92.2 0.59 5759 2226 1923 5.9 37 4.3
Rice 11.1 0.120.13 0.110.13 0.60.66 0.10.2 66 1633 3.9 1.2 0.11 1.72.0
(brown)
Teff 1.8 0.73 0.47 0.62 NA 73 2026 8 NA NA NA
Sorghum 1.32.5 0.21.1 0.61.6 0.150.39 63 24 6 NA NA NA NA
Pearl 2.22.8 0.080.16 1.31.8 0.650.84 5370 2032 13.3 NA NA NA NA
millet
Finger 0.590.69 0.140.19 0.200.24 0.060.08 55.763.4 14.438.6 18 NA NA NA NA
millet
Foxtail 0.46 0.10 0.15 0.04 5269 11.232 NA NA NA NA NA
millet
Proso NA NA 0.66 0.08 6168 17.232.6 NA NA NA NA NA
millet
Corn 1.03.0 0.20.5 0.10.4 0.91.9 0.10.3 6478 24 13.4 5.86.6 0.1 2.1
Oats 1.4 0.05 0.09 0.64 0.19 4361 1627 9.6 7.7 3.96.8 6.012.9
Rye 3.2 0.08 0.10 1.9 0.4 69 2431 14.6 8.5 1.92.9 1.5
Canary 1.75 0.06 0.07 0.83 NA 61 17.2 6.0 NA 1.12.3 NA
Triticale NA 0.23 3.1 0.21.9 NA 5970 23 1316 NA 0.51.0 1.72.5
Quinoa 2.9 0.02 0.02 NA NA 63 1112 9.6 NA NA NA

transitory starch, as it is degraded to maltose, glucose, hexose
phosphates, and triose phosphates at night. This periodical
starch degradation involves two major steps: initiation, which
involves the activation of starch, and starch digestion. Transient
starch degradation is mainly attributed to b-amylases, and the
product of its action is transported back to the cytosol, where it
undergoes further modifications.
Some of the starch degradation products are converted to
sucrose, the major transport sugar. Sucrose through symplastic
or apoplastic transport enters the phloem, through which it is
mobilized to the storage tissues, where symplastic or apoplastic
uptake into sink cells takes place. The transported sucrose is
further converted to substrates that give rise to the diverse carbo-
hydrates present in grains of different crops as described earlier.
The hexose phosphate pool also plays an important role in starch
and sucrose utilization by plants. In the glycolytic pathway,
which acts in reverse of the triose phosphate synthetic pathway
during photosynthesis, starch and sucrose are oxidized to pro-
vide energy and hexose phosphates that on phosphorylation
contribute to the hexose phosphate pool. In addition, the glyco-
lytic reactions provide several intermediates that contribute to
the synthesis of proteins, lipids, and organic acids (Figure 2).
These compounds are important grain components and are
described in other articles.
The hexose phosphate pool is an important converging
point for both carbohydrate synthesis and degradation and
thus is considered to play an important role in carbohydrate
metabolism in plants. The main components, glucose 1-phos-
phate, glucose 6-phosphate, and fructose 6-phosphate, are
readily interconvertible when present in the same system. How-
ever, in plants, the hexose phosphates are compartmentalized
to the plastids and cytosol. In photosynthetic tissues, the plas-
tidial and cytosolic hexose phosphate pools are independent of
each other, and their exchange is often restricted. Therefore, the
regulation of carbohydrate metabolism in plants is more com-
plex than in other organisms, where such subcellular compart-
mentalization of hexose phosphates is not present.
Sucrose
Sucrose is the most abundant disaccharide and the major
product of photosynthesis. It is a nonreducing sugar with
limited chemical reactivity and thus is utilized as a transport
and storage molecule in most plants. Sucrose synthesis is pre-
dominant in leaves, but the ability to synthesize sucrose is
fairly widespread among plant cells. Sucrose is synthesized in
the cytosol by the consecutive activities of two enzymes:
sucrose-phosphate synthase (SPS) and sucrose phosphatase
(SPP). SPS catalyzes the reversible transfer of glucosyl units
from UDP-glucose to fructose 6-phosphate, synthesizing
sucrose 6-phosphate that is dephosphorylated by SPP to yield
sucrose in an irreversible reaction. SPS is the key regulating step
of sucrose biosynthesis. In plants, SPS can be activated alloste-
rically by glucose 6-phosphate and inhibited by inorganic
phosphate (Pi). In addition, SPS also contains several regula-
tory phosphorylation sites. Sucrose synthase (SucS), present in
both soluble and membrane-bound forms, is responsible for
catalyzing both the synthesis and degradation of sucrose. How-
ever, sucrose degradation is mainly attributed to SucS in sink
tissues participating in sucrose degradation.
The translocation of synthesized sucrose from source
to sink tissues is essential for proper plant growth and devel-
opment. Sucrose can be translocated either symplastically
(through plasmodesmata) or apoplastically (through apoplast-
s with the help of sucrose transporters). In sink tissues, sucrose
can be stored inside cell vacuoles through transporters located
 166
CARBOHYDRATES | Carbohydrate Metabolism
Table 3 Carbohydrate composition of some grain legumes

Soluble carbohydrates (%) Starch (%) Nonstarch complex carbohydrates (%)

Total soluble Total Dietary

Crop sugars Sucrose Raffinose Stachyose Verbascose galactosides Total Amylose Polysaccharides fiber Hemicellulose Cellulose

Common 2.09.6 1.63.9 0.22.5 0.23.9 0.11.8 0.48.0 5151.9 22.136.0 6.420.4 11.227.5 0.55.6 3.213.1
bean
Peas 3.513.8 0.95.4 0.42.3 0.34.2 04.3 2.39.6 24.757.4 23.533.1 NA 16.121.6 0.912.4 0.913.3
Lentils 3.39.5 1.13.0 0.10.8 1.14.0 06.4 1.87.5 40.157.4 20.745.5 6.914.7 11.021.4 1.215.7 3.514.8
Chickpeas 4.614.2 2.86.9 00.3 0.42.0 trace0.4 2.07.6 43.059.0 31.845.8 5.535.4 8.224.0 0.616.0 1.113.7
Faba beans 2.28.5 0.13.8 0.11.5 0.21.6 1.12.4 1.04.5 39.247.2 22.035.0 17.5 17.123.8 1.68.9 8.314.3
Mung bean 3.97.2 0.32.0 0.32.6 1.22.8 1.73.8 NA 37.053.6 13.835.0 NA 15.7 0.39.1 2.54.6
Black gram 3.07.1 0.71.5 0.01.3 0.93.0 3.43.5 NA 32.247.9 43.9 NA 22.4 10.7 5.0
Red gram 3.510.2 2.7 1.01.1 2.73.0 4.04.1 NA 40.448.2 38.6 NA 18.7 10.1 7.3

Black gram: Vigna mungo; red gram: Cajanus cajan; faba bean: Vicia faba.
 CARBOHYDRATES | Carbohydrate Metabolism 167

Cell-wall carbohydrates, Starch

e.g., cellulose, -glucan, xylans

Light CO2 H2O

Glucose Hexose-phosphate pool
Sucrose ADP-glucose
Fructose Glucose-1-phosphate
Glucose-6-phosphate Photosynthesis (Calvin cycle
Raffinose reduction phase)
Fructose-6-phosphate
Fructan Glyceraldehyde-3-phosphate

Ribulose-1,5-
Purines
bisphosphate
Ribulose5-bisphosphate Triose-phosphate pool Tryglycerides
Nucleic acids 3-PGA, GAP, DHAP
Amino acids
Pentose phosphate pathway

Aspartic acid Citric acid cycle Glycolysis

pay-off phase Purines

Other amino acids, proteins, Tryglycerides

NADH ATP
nucleic acids, alkaloids
FADH2 Amino acids

Other amino acids, proteins Glutamic acid NADH Purines

Tryglycerides
Delta-aminolevulnic acid Oxidative
phosphorylation Amino acids

Chlorophyll, NAD+
Cytochrome, FAD
phytochrome ATP

Figure 2 A schematic overview of the carbohydrate metabolism and its interrelationship with other biochemical pathways to show the
production of major plant cell constituents.

at tonoplasts or can be hydrolyzed by invertase providing
glucose and fructose to the cell. The translocation of sucrose
from source to sink tissues is very active during grain develop-
ment when storage carbohydrates are synthesized in the grains.
A boost in sucrose concentration in seeds is observed during
seed germination when stored carbohydrates are broken down
to provide energy to the germinating embryo.
Alpha-Galactosides
Alpha-galactosides are low-molecular-weight, nonreducing oli-
gosaccharides and rank next to sucrose among abundant soluble
sugars. These sugars constitute about 618% of the dry weight of
mature legume seeds. Alpha-galactosides are sucrosylgalacto-
sides and are characterized by the presence of an a(1! 6) link-
age between the galactosyl residue and the C-6 of the glucose
moiety of sucrose. Alpha-galactosides can further be classified
into two groups. Raffinose family oligosaccharides (RFOs) con-
stitute the first group, and the first member of this group, raffi-
nose (degree of polymerization (DP) 3), is the main RFO in
most monocotyledon seeds, while its higher homologues, sta-
chyose (DP 4), verbascose (DP 5), and ajugose (DP 6),
accumulate predominantly in seeds of dicotyledons. Higher
members like ajugose are generally found in trace quantities in
seeds. Some alternative RFOs such as lychnose, stellariose,
mediose, and manninotriose have been recently reported from
Caryophyllaceae and Lamiaceae plants (Figure 1). The second
group of a-galactosides includes galactosyl cyclitols. Ciceritol is
the most common of the galactosyl cyclitols and was first
reported from chickpea (Cicer arietinum), followed by lentil
(Lens culinaris).
RFO biosynthesis occurs in the cytosol and begins with the
formation of galactinol from myo-inositol and UDP-galactose in
a reaction catalyzed by galactinol synthase. In the next step,
raffinose synthase supports galactinol to donate its galactosyl
unit to sucrose yielding raffinose. Raffinose and stachyose,
together with galactinol, synthesize stachyose and verbascose
in two separate reactions catalyzed by stachyose synthase and
verbascose synthase, respectively. In a galactinol-independent
pathway, already present RFOs react with each other to
synthesize higher members of RFOs. Galactan/galactan
galactosyltransferase is the sole enzyme catalyzing the reaction.
RFOs participate in plant physiological processes like seed
desiccation tolerance, seed germination, photosynthate trans-
location (mostly in Cucurbitaceae, Verbenaceae, Lamiaceae,
Oleaceae, and Scrophulariaceae families), and stress tolerance.
However, in the human diet, food rich in RFOs causes stomach
discomfort, flatulence, and diarrhea in humans and mono-
gastric animals, as they lack a-galactosidase, a hydrolyzing
enzyme needed for RFO breakdown. RFOs when consumed
in lower concentrations are considered as prebiotic and sup-
port the growth of beneficial intestinal microflora.
168 CARBOHYDRATES | Carbohydrate Metabolism

Fructans in their physiological role in the translocation and storage of

carbohydrates.
In contrast to tropical and subtropical grasses, which accumu-
Sugar alcohols are synthesized from hexoses or hexose phos-
late sucrose and starch as storage carbohydrates, grasses of
phates, through the consecutive action of reductases and phos-
temperate and cool regions mainly store fructans, the fructose
phatases. Mannitol biosynthesis begins in the cytosol with the
polymer. Fructans serve as soluble storage carbohydrates in
formation of mannose 6-phosphate from fructose 6-phosphate
about 1215% of all flowering plants belonging to the Aster-
in a phosphomannose isomerase-catalyzed reversible reaction.
ales, Poales, and Liliales. Fructans can be found as long-term
Mannose 6-phosphate is first reduced to mannitol 1-phosphate
reserves such as in bulbs and tubers or as temporary stocks in
and converted to mannitol in reactions catalyzed by mannose
leaves, stems, and seeds. In cereals, fructans can be found in
6-phosphate reductase and mannose L-phosphate phosphatase,
young internodes, from which fructan-derived carbohydrates
respectively. Sorbitol is similarly reduced and dephosphorylated
are mobilized to kernels during grain filling, especially during
from glucose 6-phosphate by the action of a glucose 6-
challenging environmental conditions. In addition, fructans
phosphate reductase and a sorbitol (glucitol) 6-phosphate
contribute to regulating osmotic balance, maintaining sink
phosphatase.
strength, and providing tolerance against cold and drought
stresses. Fructans are also designated as low-calorie sweeteners,
fat replacers, and prebiotics (fructans with low DP) in the Starch
human diet.
Starch is the most abundant storage polysaccharide in seeds,
Fructans, on the basis of glycosidic linkage, are categorized
fruits, tubers, and roots of higher plants. Cereal and tuber starch
into five groups: (a) inulin having b(2 ! 1) linkage, (b) levan/
contribute about 50% of the calories in the human diet. Besides
phlein having b(2 ! 6) linkage, (c) graminin (having inulin or
being an important energy source in the human diet and ani-
levan backbone with  1 short branch), (d) inulin neoseries
mal feed, its amenability to modification led to many nonfood
(like inulin but one glucose unit between two fructose moie-
applications including paper, pulp, textile production, cos-
ties), and (e) levan neoseries (like levan but one glucose unit
metics, packaging, biodegradable plastic films, construction,
between two fructose moieties) (Figure 1).
and mining. Since starch is a renewable and biodegradable
Fructan biosynthesis is sensitive to photosynthetic demand
resource, it has received increased interest for the production
and sink sucrose concentration. It is synthesized in vacuoles
of environmentally friendly biomaterials. However, environ-
by fructosyltransferase enzymes utilizing sucrose as the primary
mental concerns and costs associated with chemical modifica-
substrate. As the first step of fructan biosynthesis, a fructose
tions are major constraints to completely utilizing starch as an
unit from one sucrose molecule is transferred to another synthe-
industrial raw material.
sizing b(2! 1) or b(2! 6) linkage in sucrose/sucrose
Starch is a complex macromolecule composed of two
1-fructosyltransferase (1-SST)- or sucrose/fructan 6-fructosyl-
glucan polymers, an essentially linear-chain amylose and a
transferase (6-SFT)-catalyzed reactions yielding 1-kestose (inulin
highly branched amylopectin. Starch is normally made up
type) or 6-kestose (levan type), respectively. Some plant species
of one-quarter amylose, with the remaining three-quarters
also synthesize 6G-kestose by transferring a second fructosyl unit
being amylopectin. The two components are arranged in a
of 1-kestose to the glucose molecule of sucrose in a reaction
three-dimensional, semicrystalline structure the starch granule.
catalyzed by fructan/fructan 6G-fructosyltransferase (6G-FFT).
Starches from different sources differ in their overall properties
These trisaccharides, 1-kestose, 6-kestose, and 6G-kestose,
due to differences in granule size distribution and shape, amy-
serve as precursors for higher members of the fructan family.
lose and lipid contents, distribution of chain length in amylo-
Elongation of the inulin- or levan-type backbone is mediated
pectin, phosphorylation, and crystallinity. For example, native
by fructan/fructan 1-fructosyltransferase (1-FFT; uses fructosyl
starches from pulse crops generally have a higher percentage of
oligosaccharides as fructosyl unit donor) or 6-SFT (uses both
amylose as compared with cereal starches (Tables 2 and 3). Most
sucrose and fructosyl oligosaccharides as fructosyl unit donor),
of the gelling and pasting behavior of extracted starch is deter-
respectively.
mined by the amylose/amylopectin ratio, amylopectin structure,
and starch granule size distribution. Genetically altered starches
derived from mutagenesis and breeding programs have already
Sugar Alcohols
been commercialized.
Sugar alcohols are primary photosynthetic products that accu- The biosynthesis of starch is a complex biochemical process.
mulate temporarily in leaves during the light period and trans- The basic biochemical mechanism is the same for transitory
located to other plant organs at night. As compared with the starch synthesis in the chloroplasts in green organs or storage
corresponding sugar, sugar alcohols have an additional hydroxyl starch in amyloplasts of storage organs. Starch biosynthesis can
group and therefore are designated as polyols, polyalcohols, or be divided into four basic steps: (i) availability of substrates for
polyhydric alcohols. Mannitol and sorbitol (or glucitol) are the starch synthesis, (ii) priming of starch granules, (iii) chain
main sugar alcohols that have been studied in plants. These initiation and elongation, and (iv) introduction of branches.
compounds are rare in monocots but contribute significantly In chloroplasts, when photosynthetic carbon fixation is greater
to transported and stored carbon in some horticultural plants, than the amount of sucrose being exported, triose phosphates
such as members of the Rosaceae, Rubiaceae, and Plantaginaceae are converted to fructose 1,6-bisphosphate and become part of
families. Sugar alcohols have been implicated in abiotic stress the hexose phosphate pool by undergoing a reaction with
tolerance. In comparison with sucrose, sugar alcohols are more fructose 1,6-bisphosphatase. Its further processing by phospho-
metabolically sequestered, and this has important implications glucoisomerase and phosphoglucomutase results in the

formation of glucose 1-phosphate. In storage tissues, sucrose is
converted into glucose 6-phosphate in the cytosolic region
from where it is imported into the amyloplasts through a hex-
ose phosphate/Pi antiporter. The glucose 6-phosphate is con-
verted to glucose 1-phosphate, which acts as a substrate for
AGPase, to synthesize ADP-glucose as in chloroplasts and act
as the primary building block required for chain elongation to
initiate starch synthesis. AGPase is an allosterically regulated
enzyme, whose activity is generally stimulated by 3-PGA and
inhibited by Pi. This allosteric regulation is further influenced
by plant species, plastid type, and subcellular location.
Studies of starch mutants have so far produced a general
consensus that AGPase, soluble starch synthases (SSs), starch-
branching enzymes (SBEs), starch-debranching enzymes
(DBEs; pullulanase and isoamylase), and possibly dispropor-
tionating enzyme (D-enzyme) catalyze the final steps leading to
amylopectin synthesis. The five major classes of SS include
GBSSI, SSI, SSII, SSIII, and SSIV, of which all but SSIV have
been shown to have distinct roles in determining starch com-
position and structure. Granule-bound starch synthase 1
(GBSSI) is the only enzyme committed to amylose synthesis
with limited contribution toward amylopectin as indicated in
some recent reports. Since several of the enzymes exist in
different isoforms, some of which vary in their subcellular
distribution, enzyme specificity, temporal activity, and interac-
tion with other enzymes; the starch biosynthesis pathway is
complex. Hilum, also known as starch granule core, indicates
the presence of a priming mechanism for starch granule for-
mation. The priming mechanism for starch glucan polymeri-
zation in plants is unclear, but it likely involves short-chain
maltodextrins that are extended and branched by the action of
GBSSI, SBEI, and SSI to form a molecule with an amylopectin-
like backbone. Glycogenin-like self-glycosylating protein has
also been proposed. The primary glucan polymer is further
polymerized through cycles of chain extensions and branching
catalyzed mainly by SSI and SBEII. DBEs have been proposed
to play a role in starch biosynthesis by trimming the ends of the
water-soluble preamylopectin molecules to produce amylopec-
tin that can be crystallized and packaged into granules. Two
distinct DBE families (isoamylase and R-enzyme/pullulanase/
dextrinase) with substrate specificity have been identified in
plants. DBEs are also suggested to degrade water-soluble glu-
cans synthesized by starch synthases and starch-branching
enzymes to prevent accumulation of phytoglycogen, which
cannot form granules. Observations made in Chlamydomonas
suggest that the D-enzyme is also involved in starch synthesis
and granule formation in addition to starch degradation.
The polymerization of amylose is initiated from soluble
maltooligosaccharides that serve as primers for GBSSI. Starch
biosynthesis can be visualized as a wavelike formation, where
high and low nodes result in the formation of regions of
different crystallinities visible as growth rings (e.g., potato
starch). Many reports have indicated that the presence of starch
biosynthetic enzymes as protein complexes and positioning of
SS, SBE, and DBE in these complexes influence amylopectin
architecture. Amylopectin chains distributed in polymodal pat-
terns form clusters with architectural specificity. These chains
are designated as A-chains (DP 612), the outermost chains;
B-chains, named B1, B2, and B3 depending on their degree of
polymerization (DP 1324 up to 50), which support A-chains
CARBOHYDRATES | Carbohydrate Metabolism 169
or other B-chains; and the central C-chain. The backbone of
starch, that is, the C-chain from which B-chains are connected
within a granule, is oriented toward the center or hilum of the
granule. Amylopectin glucan chain length distribution influ-
ences functional properties like gelatinization, retrogradation,
and pasting properties of starch.
Cellulose
Cellulose is the major component of cell walls and accounts for
2830% of dry matter in forage grasses and 4245% of wood.
This extracellular polysaccharide-based outer matrix of the
plant cell is complex and dynamic and has enormous strength.
It is the most abundant renewable biomass synthesized on
Earth and has since the early days of human civilization been
used for fuel, timber, fiber, and forage. Today, cellulose has
additional uses as a raw material for the production of paper,
pulp, and many chemicals. Cellulose has a paracrystalline struc-
ture formed by b-(1 ! 4)-D-glucan chains, formed individually
and crystallized to form cellulose microfibrils (an average thick-
ness of 36 chains through inter- and intramolecular hydrogen
bonds and van der Waals forces). The chains contain a few
thousand glucose units giving an individual length of 23 mm,
but the chains start and end at different points of the microfi-
bril, thus extending the length of the microfibril to hundreds of
micrometers. The length, angle, and crystallinity of these micro-
fibrils influence plant cell wall strength.
In higher plants, cellulose synthesis occurs in a plasma
membrane-bound complex known as rosette TC. The structure
contains six subunits, each composed of six glucan (cellulose)
synthase molecules that are symmetrically arranged into a
rosette. The subunits of the complex simultaneously synthesize
36 1,4-b-D-glucan chains that are bundled into a microfibril
emerging from the rosette TC. The basic biochemical under-
standing of cellulose synthase (CesA) has mainly been obtained
from studies of the bacterium, Acetobacter xylinum. In plants, the
number of CesA genes varies depending upon the species.
CesAs categorized under the glycosyltransferase-2 (GT-2) super-
family with eight-transmembrane topology consist of at least
two subunits, one with a catalytic role and the other with a
regulatory role. The catalytic subunit of CesA is a progressive
glycosyltransferase that binds the substrate, UDP-glucose,
derived from sucrose by the action of a membrane-bound
SucS. Recent studies proposed that these subunits are placed
in the loop between transmembrane domains 2 and 3 with a D,
D,D,QXXRW motif. The present model of cellulose synthesis
postulates that the first two conserved aspartic acid residues
interact with UDP, the third aspartic acid residue forms the
catalytic base for the glucan extension, and the last QXXRW
residues form a binding site for the terminal glucan parts of the
chain. At the initiation of polymerization, two UDP-glucose
molecules are present in the substrate-binding site, and as the
chain elongates, glucose is added to the nonreducing end of the
chain. In addition to Ces genes, plants have a number of cellu-
lose synthase-like (CSL) genes that share homology with Ces
and belong to the GT-2 superfamily. CSLs are grouped into ten
families designated from A to J. Many of these are associated
with b-glycan polymer synthesis, for example, CSLF for
b-(1 ! 3;1 ! 4) glucans.
170 CARBOHYDRATES | Carbohydrate Metabolism
In addition to Ces, membrane-anchored endo-1,4-
b-glucanases may also play a role in cellulose biosynthesis,
based on studies with the Arabidopsis KORRIGAN mutant and
the cellulose-producing bacterium Agrobacterium tumefaciens.
Other Cell Wall Carbohydrates
Other important polysaccharide components of the cell wall
include cross-linking glycans, which are hydrogen-bonded to
cellulose microfibrils. These glycans coat the cellulose micro-
fibrils and link two or more cellulose microfibrils to form a
network in the primary cell walls of all plants. The two major
classes of the cross-linking glycans are xyloglucans (XyGs)
and glucuronoarabinoxylans (GAXs), of which XyGs are pre-
sent in the cell walls of most dicots and almost half of all
monocot species. The XyGs are made up of linear chains of
1,4-b-D-glucan with numerous a-D-Xyl units linked to the O6
position of the glucan units. In some cases, the xylosyl units
are further substituted for by arabinose or galactosyl units. In
addition to the two xyloglucans, some members of the Poa-
ceae family, which includes the majority of cereal crops, such
as wheat and barley, contain a third cross-linking glycan, b-
glucan. This is a linear polymer that consists of 3-D-
glucopyranosyl monomers joined by both (1 ! 3) and
(1 ! 4) linkages, normally in a 2:1 ratio. In comparison
with other cereals, barley and oat grains are enriched with
b-glucans. The b-1,4-linked molecules cellotriose (DP3) and
cellotetraose (DP4) are randomly joined by b-1,3 bonds,
which results in the formation of kinks in the molecule. The
DP3/DP4 ratio affects polymer solubility. The highest content
of soluble (1,3;1,4)-b-glucan is associated with DP3/DP4
ratios in the 1.52.5 range. Both the ratio and concentration
of b-1,4-linked molecules in the grain cell wall, especially in
oat and barley, influence their end use in food, feed, or
industrial applications.
H2O
-amylase
Starch
Maltose
Glucose
H2O GA
Scutellum
Coleoptile
Embryo
Radicle
Coleorrhiza
Figure 3 A generalized diagram of a germinating cereal grain to show the various parts of a grain and the major biochemical activities during
germination.
Arabinoxylans and b-glucans affect the quality and utiliza-
tion of cereal grains due to their presence in the endosperm cell
walls. The predominant component of starchy endosperm cell
walls in barley, oats, and maize is b-glucan, whereas arabinox-
ylans are the major component found in rye and wheat. These
compounds have a significant impact on the utilization of the
grains for both food and animal feed. In wheat flour, pentosans
absorb water and significantly affect the quality of baked prod-
ucts. The cell wall carbohydrates are not completely digested in
the stomach or small intestine of humans or other monogastric
animals but are broken down by bacteria in the large intestine.
They tend to reduce the absorption of mono- and disaccharides
in the gut by raising the viscosity of the gut contents, thus
decreasing the levels of blood glucose and insulin. b-Glucans
have been shown to significantly reduce blood cholesterol
levels, when present in the diet at high concentrations. In addi-
tion, they have also been shown to have antibacterial and anti-
viral effects and accelerate wound healing. In animal feed, both
glucuronoarabinoxylans and b-glucans have antinutritional
effects, as these may decrease the absorption of nutrients due
to their high viscosity and indigestibility.
Cell wall polysaccharides are synthesized from sucrose
through the action of SucS, UDP-glucose dehydrogenase, UDP-
glucuronate decarboxylase, xylan synthase, and arabinosyltrans-
ferase that sequentially give rise to UDP-glucose, UDP-glucuro-
nate, UDP-xylose, 1,4-b-xylan, and arabinoxylan, respectively.
The action of three epimerases further converts UDP-glucose to
UDP-galactose, UDP-glucuronate to UDP-galacturonate, and
UDP-xylose to UDP-arabinose.
Carbohydrate Breakdown
Germination is initiated when the dry seed imbibes water,
and the process is completed with the elongation of the
embryonic axis (Figure 3). One of the first signs of metabolic
Husk
Pericarp
Testa
Aleurone
Endosperm
GA receptor
GA
GAMyb
Aleurone cell
-amylase
Scutellum
Shoot
Root

activity in a germinating seed is the activation of enzymes
that hydrolyze storage reserves. This process has been well
characterized in cereal grains (Figure 3). A group of plant
growth regulators known as the gibberellins (GAs) are the
first to be synthesized in the scutellum of the embryo. The
GAs diffuse into the aleurone layer and induce the produc-
tion of hydrolytic enzymes such as a-amylases active on the
starch granules in the endosperm. The GA-inducible R2R3-
MYB transcription factor known as GAMYB was first identi-
fied in the cereal aleurone. It has been proposed that a GA
receptor present on the aleurone membrane binds to GA,
which induces GAMYB activation, which further binds to
the GA response element (GARE), 50 AACA/TA-30 , to activate
hydrolase genes including a-amylase involved in starch break-
down and mobilization for embryo growth. The hydrolysis
products are taken up by the embryo to initiate its growth.
Carbohydrates stored in the endosperm or cotyledons are the
primary source of energy in the developing embryo until it
forms leaves, and the photosynthetic process takes over to
supply the necessary energy and chemical constituents for the
developing plantlet.
Starch Breakdown
In cereal grains, starch is the major carbohydrate that is broken
down during germination. Starch degradation occurs in three
distinct phases: (1) reduction of starch granules to soluble
maltodextrins, (2) degradation and debranching of larger mal-
todextrins to glucose and glucose 1-phosphate, and finally (3)
transport of the glucose or glucose 1-phosphate from the stor-
age organs to the hexose phosphate pools of developing plant
cells for further metabolism into chemical constituents and
energy. In germinating seeds, the initial degradation of starch
granules is associated with an increase in the a-amylase activ-
ity. Intact starch granules are attacked by a-amylase resulting
in the formation of random pits on the granules. During
the a-amylase attack, a(1,4) bonds in both amylose and
amylopectin are randomly broken to release large glucan
polymers. Subsequent degradation occurs with assistance
from b-amylases, phosphorylases, maltases, and starch DBEs.
a-Amylase action on amylopectin, which contains both a(1,4)
and a(1,6) glucan linkages, results in maltose and branched
dextrins of short chain length as a-amylase cannot break the
a(1,6) linkages. b-Amylases first act only on the nonreducing
ends of the glucan polymer and hydrolyze starch into
b-maltose, which is rapidly converted into natural mixtures of
a- and b-isomers by mutarotation. Starch phosphorylases are a
group of phosphorolytic enzymes that degrade starch mole-
cules from the nonreducing end of the glucan chain. The
reaction results in the complete breakdown of amylose to
glucose 1-P units, but in amylopectin, the degradation stops
within a few glucose residues of the branch point to give
dextrins. Like plant amylases, starch phosphorylases are also
widely distributed in plants and have an important role in
starch degradation in leaves and seeds of many plants.
However, in germinating cereal grains, starch phosphorylase
activity is not detected, giving the a- and b-amylases a domi-
nant role in starch degradation. In the seeds of other species
and in leaves, starch phosphorylases also contribute toward
starch degradation.
CARBOHYDRATES | Carbohydrate Metabolism 171
The a(1,6) glucan linkages in the starch molecule are
hydrolyzed by starch DBEs, which have also been implicated
in starch synthesis as described earlier. This group of enzymes
hydrolyze a(1,6) linkages, thus producing additional end
groups for action by amylases and phosphorylases. Concerted
action of these enzymes results in the production of maltose,
glucose, or glucose 1-phosphate. Maltose rarely accumulates
during starch digestion as it is broken down to glucose by
maltase. Starch breakdown is influenced by its structure and
composition. Amylopectin fine structure and amylose both
play important roles in effecting its breakdown by metabolic
enzymes.
Raffinose Family Oligosaccharide
RFOs, accumulated in seeds by phloem loading and transport,
are rapidly utilized during the early stages of seed germination.
Alpha-galactosidase cleaves the terminal nonreducing
a(1 ! 6)-linked galactose residues of a-galactosides, thus pro-
viding readily available energy and carbon to the germinating
seed. Two types of a-galactosidases have been reported (1)
acidic (found in protein storage vacuoles (PSVs)) and (2)
alkaline (active in cytosol). Acidic a-galactosidase (activated
by the pH shift during seed imbibition) was proposed to be
responsible for the breakdown of RFOs in PSVs during the
early stages of seed development, providing the main energy
sources for seed metabolism. The resulting increase in sucrose
concentration during the transition phase, from germination
to plant growth, was attributed to the induction of expression
of alkaline a-galactosidase, resulting in the mobilization of the
remaining RFOs. The galactose unit is thereafter phosphory-
lated by a galactokinase to produce a-D-galactose 1-P, which is
further metabolized through UDP-galactose to produce either
UDP-glucose or glucose 1-P, which enters the cellular hexose
phosphate pool. Depending on the metabolic activities of the
tissues, the carbon may flow into the respiratory pathways as in
the case of germinating tissues or to the synthesis of starch,
sucrose, oligosaccharides, or other cellular components such as
glycolipids or glycoproteins (Figure 2) in young meristematic
tissues. Most of the RFO-degrading enzymes have been puri-
fied from seeds of pulse crops, and increased activities have
been shown to be associated with seed germination. However,
strategies to modulate the activities of RFO-degrading enzymes
are complicated by the presence of two distinct pathways
for the metabolism of galactose 1-P to UDP-glucose and
glucose 1-P.
Fructans
Fructans are hydrolyzed by fructan 1-exohydrolases (1-FEH) or
fructan 6-exohydrolases (6-FEH) that cleave the terminal
b(2,1)- or b(2,6)-linked fructosyl moiety, respectively. These
enzymes are also proposed to participate in fructan biosynthe-
sis by modulating the chain length and branching pattern of
the polymer. A b-fructofuranosidase from Jerusalem artichoke
tubers successively cleaves fructose units from inulin until a
mixture of fructose and the terminal sucrose unit remains.
Fructose can directly enter the respiratory pathway, whereas
sucrose must be broken down to glucose and fructose first.
172 CARBOHYDRATES | Carbohydrate Metabolism
Sucrose
Invertase and SS are the two enzymes capable of cleaving
sucrose present in plants. Invertases catalyze the irreversible
hydrolysis of sucrose to free glucose and fructose. Based on
optimum pH of their activity, invertases are grouped into acidic
(optimum pH 4.05.5) and alkaline/neutral types (optimum
pH 7.08.0). Acidic invertases are mainly found in cytosol and
occasionally in cell walls, whereas alkaline/neutral types are
abundant in vacuoles and cell walls. Alkaline/neutral invertases
have recently been reported in chloroplasts and mitochondria.
Alkaline invertases are sucrose-specific, while neutral invertase
from carrot can also mediate raffinose and stachyose hydrolys-
es. The cell wall invertases hydrolyze the incoming translocated
sucrose into glucose and fructose molecules.
SucS catalyzes the reversible reaction converting sucrose
and UDP to fructose and UDP-glucose. Fructose is available
for respiration and the UDP-glucose can enter the hexose
phosphate pool for further metabolic processes. SS, compart-
mentalized in the cytosol, is the main enzyme that degrades
sucrose in starch storage organs such as developing seeds or in
rapidly growing tissues that are converting sucrose to cell wall
structural polysaccharides. In slow-growing and mature cells,
invertase is the major enzyme hydrolyzing sucrose, which pro-
vides substrates for respiration.
Cell Wall Polysaccharides
The cell wall polysaccharides have a physiologically different
role in seed germination. The process of germination and
seedling development requires the participation of several
endo- and exohydrolases that degrade or modify the cell wall
structure. During germination, (1,3)- and (1,4)-b-D-glucan
endohydrolases hydrolyze the (1,4) linkages adjacent to the
(1,3)-b-D-glucosyl residues to release large oligosaccharides.
The resulting oligosaccharides are further hydrolyzed by
broad-specificity exohydrolases to produce glucose, which can
subsequently be used in numerous ways (Figure 2). Some
other cell wall polysaccharides, such as arabinoxylans, can be
degraded to oligosaccharides that subsequently follow the oli-
gosaccharide degradation pathway as described earlier.
Concluding Remarks
In most grains, carbohydrates are the major components that
provide the energy needed during seed germination and early
seedling development. Carbohydrates also play a major role in
determining the quality of the grain and its subsequent use by
human beings. Carbohydrate metabolism in plants differs from
most other organisms as it occurs in two distinct cellular com-
partments, the plastid and the cytosol. Most of the biosynthetic
activities take place in the plastids, chloroplasts in leaves, and
amyloplasts in the storage organs. The carbohydrate metabo-
lism in the cytosol communicates with that of the plastids
through metabolic carriers in the plastid envelope. The station-
ary nature of the plant forces it to withstand changes in the
external environment, and therefore, plants have developed
inherent flexibility in their metabolism to cope with both biotic
and abiotic stresses. It is often found that a single reaction in the
carbohydrate metabolic pathways is catalyzed by multiple iso-
forms of an enzyme. Differences in the temporal and spatial
expression of genes encoding these isoforms present several
challenges to modifying the carbohydrate metabolic pathways
by genetic engineering. To date, it has been possible to alter the
expression of genes encoding starch biosynthetic enzymes to
produce crops with modified grain carbohydrates that have
health benefits for human beings and/or grain processing
advantages. Some advances have also been made in changing
the cell compartment and/or tissue in which a specific polysac-
charide is stored. As knowledge of the carbohydrate metabolic
pathway expands, novel strategies to alter the carbohydrate
biosynthetic or degradation pathways will emerge, which can
be utilized to develop crop cultivars with novel products in the
grain. These new crops will diversify the uses of the grain to
benefit both the producers and consumers.
Learning Outcomes
Hexose phosphate pool serves as a central component of
carbohydrate metabolism.
Sucrose and raffinose family oligosaccharides maintain
sourcesink relationship by photosynthate translocation.
Starch, the most abundant storage polysaccharides, in
plants exists as a temporary/transient form in leaves and a
stable storage form in grains and tubers.
Cellulose and glycans are important for structural integrity
of cell wall.
Complex and highly regulated enzymatic machinery is
involved in structural and storage carbohydrate biosynthe-
sis and metabolism. Its key modulator steps with possibil-
ities of genetic alterations provide ample opportunities of
crop improvement for food and industrial usage.
Exercises for Revision
Question 1: Differentiate oligosaccharide from polysaccha-
ride with example. Which group of food grains has the
highest concentration of oligosaccharides?
Question 2: What are dietary fibers and what role do they
play in human nutrition?
Question 3: Describe two major metabolite pools of the
reductive pentose phosphate pathway with their major
functions inside the plant system.
Question 4: Describe the major steps of starch biosynthesis,
with emphasis on the enzymes involved in each step.
Question 5: In a germinating seed, what is the major starch-
metabolizing enzyme and how is it activated?
Question 6: How do tropical grasses differ from temperate
grasses in carbohydrate storage?
Exercises for Readers to Explore the Topic Further
How do plantenvironment interactions affect carbohy-
drate metabolism in plants?

What are the mechanisms attributed for stress tolerance by
sugars?
How does genetic modification of one biosynthetic path-
way affect the hexose phosphate pool and the biosynthesis
of other carbohydrates?
What are the critical limits (minimum and maximum) of
carbohydrates concentrations required for normal plant
growth and development?
How are sugars involved in the genetic regulation of meta-
bolic pathways?
Acknowledgments
We gratefully acknowledge the following for supporting our
research: Canada Research Chairs, Natural Sciences and Engi-
neering Research Council, Western Grains Research Founda-
tion, Saskatchewan Winter Cereals Development Commission,
Winter Cereals Manitoba Inc., Agriculture and Agri-Food
Canada, Saskatchewan Pulse Growers, and Saskatchewan
Agriculture Development Fund.
See also: Carbohydrates: Cereals: Chemistry and Physicochemistry
of Non-starchy Polysaccharides; Grains other than Cereals: Non-starch
Polysaccharides; Starch: Analysis of Quality; Starch: Chemistry; Starch:
Synthesis; Fats: Lipid Chemistry; Processing of Grains: Starch:
Modification; Starch: Uses of Native Starch; Proteins: Protein
Synthesis and Deposition; The Basics: The Grains that Feed the
World.
Further Reading
Aoki N, Hirose T, and Furbank RT (2012) Sucrose transport in higher plants: From
source to sink. In: Eaton-Rye JJ, Tripathy BC, and Sharkey TD (eds.)
Photosynthesis: Plastid Biology, Energy Conversion and Carbon Assimilation.
Advances in Photosynthesis and Respiration, vol. 34, pp. 703729. Berlin: Springer
Science Business Media B.V.
Blochl A, Peterbauer T, Hofmann J, and Richter A (2008) Enzymatic breakdown of
raffinose oligosaccharides in pea seeds. Planta 228: 99110.
CARBOHYDRATES | Carbohydrate Metabolism 173
Buchanan BB, Gruissem W, and Jones RL (eds.) (2002) Biochemistry and Molecular
Biology of Plants. Rockville, MD: American Society of Plant Physiologists.
Burton RA, Collins HM, Kibble NAJ, et al. (2010) Over-expression of specific HvCslF
cellulose synthase-like genes in transgenic barley increases the levels of cell wall
(1,3,1,4)-b-D-glucans and alters their fine structure. Plant Biotechnology Journal
9: 119.
Chibbar RN, Ambigaipalan P, and Hoover R (2010) Molecular diversity in pulse seed
starch and complex carbohydrates and its role in human nutrition and health. Cereal
Chemistry 87: 342352.
Chibbar RN and Baga M (2003) Genetic modification of primary metabolism
Carbohydrates. In: Thomas B, Murphy DJ, and Murray BJ (eds.) Encyclopedia of
Applied Plant Sciences, pp. 449459. Oxford: Elsevier.
Cui SW and Wang Q (2009) Cell wall polysaccharides in cereals: Chemical structures
and functional properties. Structural Chemistry 20: 291297.
Cummings JH and Stephen AM (2007) Carbohydrate terminology and classification.
European Journal of Clinical Nutrition 61: S5S18.
Edwin JR, Emilie AR, Veronique A, Lailiang C, and Robert T (2009) Phloem loading
strategies in three plant species that transport sugar alcohols. Plant Physiology
149: 16011608.
Hedley CL (2001) Carbohydrates in Grain Legume Seeds: Improving Nutritional Quality
and Agronomic Characteristics. Wallingford: CABI Publishing.
Hrmova M and Fincher GB (2001) Structure-function relationships of b-D-glucan endo-
and exohydrolases from higher plants. Plant Molecular Biology 47: 7391.
Jaiswal S, Ganeshan S, Baga M, and Chibbar RN (2010) In Planta modification of starch
quantity and quality. In: Singh BP (ed.) Industrial Crops and Uses, pp. 236258.
Wallingford, UK: CAB International.
Lunn JE (2008) Sucrose metabolism. In: Encyclopedia of Life Sciences. Chichester:
Wileyhttp://dx.doi.org/10.1002/9780470015902.a0021259.
Martinez-Villaluenga C, Frias J, and Vidal-Valverde C (2008) Alpha-galactosides:
Antinutritional factors or functional ingredients? Critical Reviews Food in Science
and Nutrition 48: 301316.
McFarlane HE, Doring A, and Persson S (2014) The cell biology of cellulose synthesis.
Annual Review of Plant Biology 65: 6994.
Yan D, Duermeyer L, Leoveanu C, and Nambara E (2014) The functions of the
endosperm during seed germination. Plant and Cell Physiology 55: 15211533.
Zeeman SC, Kossmann J, and Smith AM (2010) Starch: Its metabolism, evolution and
biotechnological modification in plants. Annual Review of Plant Biology
61: 209234.
Relevant Websites
http://www.chem.qmul.ac.uk International Union of Pure and Applied Chemistry and
International Union of Biochemistry and Molecular Biology; IUPAC-IUBMB Joint
Commission on Biochemical Nomenclature (JCBN) Nomenclature of
Carbohydrates.
http://www.fao.org Includes classification and chemistry of carbohydrates,
carbohydrate metabolism in fish, factors affecting it and energy transformation.
 Starch: Chemistry
FL Stoddard, University of Helsinki, Helsinki, Finland
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Starch is a widespread polymer in the plant kingdom,
composed of chains of a-1,4 linked glucose in helices,
bridged by a-1,6 bonds.
It develops in granules within chloroplasts and amyloplasts.
Two classes of starch molecule are recognized, highly
branched amylopectin and poorly branched amylose.
Packing of amylopectin double helices confers diagnostic
x-ray diffraction patterns.
Starch granule morphology differs between families of
plants.
In the presence of heat and moisture, starch helices open
and the granules are disrupted.
Cooked, cooled starch forms a paste or gel with properties
that depend on amylose and amylopectin concentration
and structure.
Cooked starch gradually recrystallizes (retrogrades), losing
water (syneresis).
Industry requires starches with different compositions
(granule size, amylose content, and amylopectin chain
length) for different uses.
Learning Objectives
To understand the role of chain length, branching pattern,
and association in starch, as these affect economically
important industrial and food uses
Introduction
Starch is the major energy reserve of higher plants and hence a
major energy source in the diet of humans and many other
animals, including livestock animals grown for their meat and
milk and pest animals that consume grains in storage. Starch
comprises
70% of the dry matter of the endosperm of a
mature cereal grain and
16% of the fresh matter of a mature
potato tuber.
Starch may be loosely defined as a polymer of glucose that
is distinguished from other glucans by a number of key
features.
First, it is a-linked, unlike cellulose and other b-linked
glucans. The majority of these links are a-1,4 (also written as
a-(1 ! 4)), leading to chains of varying lengths, and the chains
are joined together by a-1,6 linkages. These branching points
occur in a highly ordered fashion, leading to double-helix
formation and a type of crystallinity, minimizing water-
holding capacity. This feature is in significant contrast to
glycogen, an a-(1 ! 4), (1 ! 6) mixed-link glucan found in
animals and bacteria. Ordering in glycogen is very limited, so
it holds much more water than an equivalent mass of starch.
174 Encyclopedia of Food Grains, Second Edition
It is thus not as efficient an energy store where water avail-
ability is limited, such as in a dry, dormant seed.
Starch is deposited in granules between 1 and 100 mm in
diameter within membrane-bound organelles. In the actively
photosynthetic parts of the plant, these organelles are the
chloroplasts, and in storage organs, they are the modified
chloroplasts called amyloplasts. Granule structure has a num-
ber of features that are consistent across species, while granule
size, morphology, and composition vary widely among spe-
cies. These aspects and starch composition affect the processing
properties of the starch.
The Monomer and Linkages
The building block of starch is D-glucose, in its hexagonal
pyranose ring conformation (Figure 1). Carbon 1 of this ring
is a highly reactive, aldehydic, reducing end. Most commonly,
during polymer formation in a plant, it attaches to carbon 4 of
another glucopyranose residue (Figure 2). When the mono-
mers are a-D-glucopyranose, the resulting polymer is a chain of
starch, and when they are b-D-glucopyranose, it is a chain of
cellulose.
An a-1,4 linked glucan is at its most stable in a left-handed
helix, with a complete turn taking six residues (Figure 3) and
occupying 2.1 nm. Hydrogen bonding between adjacent glu-
cose residues helps to stabilize the helix. The helix can allow a
second parallel helix to fit into it, creating a double helix. The
helical structure of starch is an important factor in its proces-
sing properties and in its enzymatic degradation during diges-
tion. Cellulose, in contrast, is highly crystalline and ribbonlike
rather than helical, and relatively few organisms have the
appropriate enzymes to hydrolyze it.
The presence of a-1,6 bonds (Figure 2) allows the develop-
ment of much larger and less soluble polymers through the
introduction of branching points connecting glucose chains of
various lengths. Starch is an excellent example of the evolu-
tionary conservation of a successful structure throughout the
plant kingdom.
Starch Polymer Types
Two classes of starch, amylose and amylopectin, have been
widely recognized, differing in chain length, branching pat-
tern, and overall mass. Both have a single reducing group per
molecule and a variable number of nonreducing ends. Most
plant starches comprise about one-fifth to one-third amylose
and the remainder amylopectin (Table 1). Plant breeders and
geneticists have considerably enhanced this range since the
beginning of the twentieth century.
Amylose is usually described as being essentially linear. A
typical amylose molecule comprises 211 chains of 200700
http://dx.doi.org/10.1016/B978-0-12-394437-5.00093-0
 CARBOHYDRATES | Starch: Chemistry 175

H 6 100  C. The single helix retains a hydrophobic channel or

4 CH2OH O lumen along its axis, into which a fatty acid, for example,
may interpolate. These amyloselipid complexes have critical
OH 5 H
effects on the processing properties of the starch, as described
H H in the succeeding text (starch-bound lipids). Amylose concen-
3
2 tration increases during the course of starch deposition in the
OH 1
OH
storage organ and, in maize, is greater in proximal than in
H OH distal grains.
Figure 1 D-Glucopyranose in its a-conformation. In the Amylopectin is a much more complex entity and is usually
b-conformation, the hydroxyl group and hydrogen attached to carbon 1 described as highly branched. The overall molecular mass is
are in the opposite orientation. of the order of 106108, making this polymer one of the largest
known in nature. Chain length distribution is multimodal and
there are three main types of chain. The A-type chain is the
outermost in the branching pattern and contains 1530 a-1,4
O O linked D-glucose residues. It is attached to a B-type chain by an
a-1,6 bond between the reducing C1 of the A-chain and a C6
1 of the B-chain. The A-chain/B-chain ratio is commonly
O
11.5:1. The B-chain has one to several A-chains attached to
6
1 4 it and is itself attached either to other B-chains or to the
O O
C-chain, which carries the reducing end of the polymer. The
O
1 B-chain is classified as B1B4, with the numeral indicating

O 4 how many crystalline lamellae, or zones of the starch granule,
it passes through. A B1-chain passes through only one lamella
O and is similar in chain length to the A-chains. A B2-chain
O passes through two such lamellae and has a chain length
of
4550 glucose residues. The B3- and B4-chains are
correspondingly longer and rarer. The currently accepted
O model, supported by a range of analytic methods, is that the
branching points are clustered within any given molecule and
Figure 2 An a-1,4 linked glucose polymer with an a-1,6 branching also coordinated in adjacent molecules, rather than random
point. (Figure 4). The model shows double helices, often of an
A-chain and a B1-chain, as the state of the outermost branches.
The lumen of a double helix is smaller than that of a single
O O
O helix and hence less likely to include guest molecules. In
potato amylopectin, phosphate groups are attached to some
O of the hydroxyl groups of the glucose residues. This is associ-
ated with greater hydrophilicity and may contribute to the
O high swelling power of potato starch, which is ten times
O that of many other starches.
O Numerous biochemical tests have been devised to evaluate
the relative proportions of amylose and amylopectin in a
O
starch. One of the most common involves the use of iodine,
which may be quantified in two ways. Amylose can bind 20%
O of its mass in iodine, whereas amylopectin binds <1%. This
O may be measured accurately, if slowly, by potentiometric titra-
O tion. In a more approximate and considerably more rapid
O
O
method, the light absorption of the solution may be measured.
Chains of triiodide ions bound by the amylose helix absorb
Figure 3 Helix of a-1,4 glucose residues, seen from above. strongly at longer wavelengths of light, 600640 nm, causing
the starchiodine solution to appear blue. Each turn of the
helix holds about two iodine atoms. Shorter chains infre-
a-1,4 linked glucose residues. The chains are connected by a- quently included in amylopectin helices absorb at a lower
1,6 bonds, leading to an overall polymer size of 10005000 wavelength, 530550 nm, and confer a tan, reddish, or pur-
glucose residues and a molecular mass of (1.68)  105. Chain plish color to the solution. Any lipid molecules in the solution
length and branch number are species-specific. Within the compete with the triiodide ions for the amylose helices and
starch granule, the amylose chain is present as a single helix. thereby reduce the apparent amylose concentration. Simulta-
This is demonstrated by the ease with which it may be leached neous measurement of absorbance at 26 wavelengths across
out of the granules by water at moderate temperatures and its the visible spectrum generally increases the accuracy of this
full release at 6080  C, whereas an amylose double helix method over the traditional measure at a single wavelength
should not be released until the temperature greatly exceeds in the 600640 nm range. High-pressure size exclusion
176 CARBOHYDRATES | Starch: Chemistry

Table 1 Typical properties of some common starches

Source Granule average diameter (mm) Amylose concentration (%) Lipid concentration (%) Protein concentration (%)

Maize (corn) 15 25 0.80 0.35

Wheat, barley, and rye 25 27 0.90 0.40
A-granules
Wheat, barley, and rye 5 24 0.90 0.40
B-granules
Rice 3 20 0.62 0.60
Oat 5 27 1.10 0.80
Amaranth 1.5 20 1.10 0.49
Quinoa 1.5 9 0.11 0.91
Canary grass 2 17 0.02 0.20
Pulses or grain legumes 25 33 0.10 0.50
Potato 3040 20 0.10 0.10
Cassava 8 17 0.10 0.10

Data from a variety of sources.

Flexible spacers in
the amorphous
lamella (~2.7 nm)

B2 chain

Amylopectin
double helices in
the crystalline
lamella (~6.3 nm)

C chain

Figure 4 Representation of the amylopectin molecule as a liquid crystal. The shaded blocks represent double helices. In the dry state (left), they are
disordered, and in the hydrated state (right), they are aligned. Adapted from Donald AM, Perry PA, and Waigh TA (2001) The impact of internal
granule structure on processing and properties. In: Barsby TL, Donald AM, and Frazier PJ (eds.) Starch: Advances in Structure and Function, pp. 4552.
Cambridge: Royal Society of Chemistry, with permission.

chromatography is a modern method for amylose determina-
tion with a reasonable balance of throughput and accuracy.
Intermediate material was a term used in maize starch
chemistry for many years, for polymers having some of the
features of amylose and others of amylopectin due to the
combined effects of different mutations in starch-synthesizing
enzymes, for example, amylopectin-type branching but with
much longer chains.
Phytoglycogen is found in certain plant mutants such as
sweet corn. It is due to a mutation in the gene for starch-
debranching enzyme or isoamylase. Its structure is very com-
parable to that of animal glycogen, and its lack of higher-order
structure, such as double helix formation, allows it to remain
largely water-soluble. This aspect limits the amount of storage
polysaccharide that can accumulate in a grain, so when it dries
down at maturity, it is shrunken and wrinkled.
Levels of Organization
The next order of structure is an alignment of the short, stiff
double helices of the amylopectin molecules. This takes place
in two ways. First, they align parallel to each other, along the
radius of the starch granule. This is what gives the polarization
of light, leading to a characteristic Maltese cross appearance
when starch granules are viewed between crossed polarizers in
a microscope. Second, the double helices pack together in an
energy-minimizing fashion (Figure 5). In some starches,

Figure 5 Packing of amylopectin double helices: left, monoclinic crystal giving A-type x-ray diffraction pattern; right, open crystal giving B-type x-ray
diffraction pattern.
generally those from tubers and stems, the packing is hexago-
nal with a lumen in the center, which leads to a consistent
pattern, termed B-type, in x-ray diffraction. In other starches,
mostly those from grains, the double helices pack much more
closely, without a lumen, leading to a different, A-type, x-ray
diffraction pattern. Starches from grain legumes and from most
roots have an intermediate diffraction pattern, termed C-type,
which implies that some of the hexagons have a lumen and
others do not. A final x-ray diffraction pattern, V-type, is given
by amylose that encloses a lipid chain. High-amylose maize
gives a B-type diffraction pattern, suggesting that amylose does
not pack as tightly as amylopectin. Some species of yam (Dios-
corea) show A-type diffraction patterns and some B-type.
Starches with the B-type pattern generally retain or take up
more water than those with an A-type pattern.
The crystalline double helices are thus aligned in a crystalline
lamella. This alignment is possible because the branches in
amylopectin are clustered, leading to an amorphous (non-
crystalline) region (Figure 4). The thickness of this amorphous/
crystalline ring is commonly 9 nm. This is commensurate with
a helix of
18 residues (6.3 nm) plus a few more residues to
join with the B-chain in the amorphous lamella. This, in turn,
accounts for the 1530 unit length of the majority of amylo-
pectin chains.
Alteration of packing can occur in the presence of gentle
hydration, leading to the model that in the hydrated starch
granule, the rodlike double helices on their flexible attach-
ments function as a liquid crystal (Figure 4).
Stacks of
16 amorphous/crystalline lamellae form semi-
crystalline layers of
140 nm in thickness. These layers are
separated by amorphous layers of comparable thickness. This
level of order was visible in early electron micrographs and
considered to be growth rings. Early research found that they
disappeared in wheat grown in continuous light, but this is not
always the case and they are generally attributed to physical
mechanisms. Most starch granules show the alternating semi-
crystalline and amorphous layers across their entire cross sec-
tion. Certain double mutants in maize, however, show clearly
biphasic growth. In ae du and ae wx lines, the core 3060% (by
volume) of the starch granule shows the typical concentric
rings, but the remaining outer mass of the granule is shapeless
and disorganized. It was proposed that there was a change in
the relative expression of the starch synthetic enzymes and in
substrate concentration.
CARBOHYDRATES | Starch: Chemistry 177
Starch Granule Morphology
Starches of wheat, rye, barley, and many of their wild relatives
in the cereal tribe Triticeae have biphasic granule initiation,
rather than biphasic growth. One major starch granule, termed
A-type, originates in each amyloplast. About 15 days into grain
filling, membrane tubules appear around the equator of the
A-granule and several subsidiary, B-type, granules form. At
grain maturity, the A-granules are approximately lenticular or
lentil-shaped, with a long diameter about twice the short dia-
meter and with a detectable equatorial groove. Mean A-granule
diameter can be as low as 15 mm in diploid wheats such as
Triticum monococcum or as large as 30 mm in some tetraploid
and hexaploid Aegilops species but is 2025 mm in cultivated
rye, barley, durum wheat, and common wheat (Table 1). The
B-granules are roughly spherical and average 56 mm in diam-
eter in most of these species. There are other differences
between A- and B-granules. B-granules lack the central cavity
or hilum found in A-granules and also in the granules of other
species. They also lack at least one granule-bound protein, and
they can have amylose contents 24% lower than A-granules.
With their greater surface area, they adsorb more water per unit
mass than A-granules. The factors determining A-granule size
and B-granule concentration or initiation are the subjects
of continuing research. Many scientists view a third population
of small B-granules as a distinct C-granule type. This bimodal
(or trimodal) granule size distribution is not found in any
other group of plant species.
In maize, starch granules are spherical when isolated from
mealy regions of the mature grain where there is space
between the granules but are polyhedral when isolated from
vitreous regions where the granules have been in close phys-
ical contact since partway through grain filling. Faceted granule
surfaces are thus attributable to restricted growing space rather
than active differences in granule development. A-type gran-
ules of wheat and barley often show indentations where other
granules have impinged on them during development. Oat
and rice have compound starch granules. Grain legume starch
granules are generally ovoid or elliptical, and root or tuber
starches are usually round to ovoid.
Zero-amylose (waxy) starches have a fairly normal gross
morphology. Granules of high-amylose starches, in contrast,
such as those of amylomaize, pure-amylose barley, or
wrinkled pea, are shapeless, fissured, and nonbirefringent.
178 CARBOHYDRATES | Starch: Chemistry
These features demonstrate the critical role of amylopectin
in determining granule crystallinity and the much less critical
role of amylose.
Starch-Bound Lipids
As mentioned in the preceding text, long amylose chains and
lipids can form complexes. The concentration of lipid in starch
is very strongly correlated with the amylose concentration of
the starch. The amyloselipid complexes resist both their own
leaching from the starch granule and also the entry of water
into the granule. This latter aspect may be the biological impor-
tance of starch-bound lipids in the quiescent or germinating
seed. Starch processing properties are also affected by the pres-
ence of bound lipid that increases gel temperature and the
temperature of maximum viscosity, decreases gel strength,
and delays staling by interfering with retrogradation. Removal
of lipids lowers gel temperature and increases peak viscosity.
Starch-bound lipids are generally either free fatty acids
(commonly linoleic) or lysophospholipids. The ratio of these
two classes differs among species in the cereals (Table 2), but
there is much less information on the pulses and pseudo-
cereals. The lipid concentration of extracted starch varies up
to tenfold depending on the extraction method.
Starch-Bound Proteins
Numerous types of protein are found in close association with
starch granules, some on the surface and others bound within.
The biological functions of some of these proteins remain to be
determined. The surface proteins tend to be smaller than the
internal proteins. Altered sequences in the tryptophan-rich
surface proteins, puroindoline-a and puroindoline-b, have
been strongly linked with difference in grain hardness, but
their causative role in this difference is controversial.
Granule-bound starch synthase (GBSS), which has a rela-
tive molecular mass of
60 kDa, is responsible for synthesiz-
ing amylose and is thus absent from the so-called waxy
mutants identified in a wide range of crops including maize,
barley, wheat, rice, pea, and cassava. Confocal laser scanning
microscopy of fluorescent-dyed proteins has shown that GBSS
is localized in concentric spheres coinciding with the amor-
phous layers of the growth rings in the starch granules of
maize, wheat, and potato. Other unidentified proteins were
shown by this method to be concentrated in the hilum of the
granule. A large, 140152 kDa form of starch branching
enzyme I (SBE) was found to be associated only with
A-granules, not B-granules, of wheat.
Table 2 Composition of starch-bound lipids of several cereals
Species Free fatty acids Lysophospholipids
Wheat, barley, and rye 10 90
Rice and oat 30 70
Millet and sorghum 45 55
Maize 60 40
Data from a variety of sources.
The total protein concentration of extracted starch is higher
in cereal grains than in roots or tubers (Table 1), although the
exact figure depends heavily on the extraction method.
Dissociation Chemistry
Gelatinization has been defined as the collapse (disruption)
of molecular order within the starch granule, manifested in
irreversible changes in properties such as granule swelling,
native crystalline melting, loss of birefringence, and starch
solubilization. Pasting is what happens after gelatinization
granule swelling, exudation of molecular components from
the granule, and, eventually, total disruption of the granules.
Starch granules in cold water can reversibly absorb 30% of
their weight in water. When starch is heated in an excess of
water, a sequence of events occurs as the kinetic energy of the
polymer increases and the hydrogen bonds rupture. At the
pasting or gelatinization temperature, between 60 and 70  C,
energy is absorbed in decrystallizing the crystalline regions of
the molecule. It also leads to an endotherm, as seen in the
differential scanning calorimeter (DSC), due to the absorption
of energy. Typical values of the heat of gelatinization or gelati-
nization enthalpy are 812 J g1 but are as high as 32 in
amylomaize. The loss of crystallinity leads to a rapid swelling
of the granule combined with a considerable loss in structure,
as seen in a rapid increase in viscosity in a pasting curve
(Figure 6). Amylose leaches from the granule but amylopectin
remains associated.
Further heating without stirring in an enclosed vessel, such
as a DSC, causes dissociation of amylose from lipid. This
process also absorbs energy and shows as an endotherm
between 102 and 110  C.
If, however, the heating is in the presence of stirring in a
nonpressurized container, such as a Brabender Visco-Amylo-
Graph or a Rapid Visco Analyzer (RVA), the viscosity of the
suspension continues to increase as the amylose molecules get
more dispersed and the granules become more porous, gelati-
nized, and swollen. Further stirring at a high temperature
results in shear thinning, where the molecular entropy is not
enhanced but granule structure is gradually further destroyed
until a new plateau is reached. This phase is known as break-
down. As the paste is cooled and stirred, viscosity reaches a
final value. The RVA takes
15 min for the process, instead of
about an hour for the Visco-Amylo-Graph. The RVA has the
significant advantage that it gives readings in SI units (cP), in
contrast to the arbitrary units of older models and the Visco-
Amylo-Graph.
In a set of rice starches of equivalent amylose content, most
RVA parameters were not significantly affected, but breakdown
was correlated negatively with the proportion of long amylo-
pectin chains and positively with the proportion of short. In a
separate study, increased firmness of cooked rice was associ-
ated with increased content of long amylopectin chains. The
short chains are types A and B1, the long B2B4. Perhaps, the
presence of more interconnections between crystalline lamel-
lae, given by a higher content of B3 and B4 chains especially,
helps to maintain structure of the gelatinized starch granule.
Furthermore, the radius of rotation of the chain increases with
its degree of polymerization, and the viscosity increases as the

Figure 6 Starch pasting profile, as determined with an RVA (Newport Scientific Pty Ltd., Warriewood, NSW, Australia). Key measurements are named
on the graph. The thin black line shows temperature profile and the thick green line shows viscosity profile.
Table 3 Classification of resistant starch
Type Description Example
1 Physically inaccessible Grain fragments
2 Native granules Uncooked banana
3 Retrograded after heat/moisture Cooled, cooked
treatment potato
4 Chemically or thermally modified Acetylated starch
cube of the radius, so small increases in chain length can lead
to large increases in viscosity.
Reassociation Chemistry
As stated earlier, the minimum energy state for an a-1,4 linked
glucan is as a double helix with another such chain. As a starch
solution cools, the chains reassociate in a process termed
retrogradation. Long amylose chains have a considerable
radius of rotation and freedom of movement in a solution
and may thus entangle with many other long chains at con-
centrations as low as 11.5%. Amylose retrogradation thus
entraps a lot of water, involves many molecules, makes a
clear hard gel, and is essentially complete when the cooked
product has cooled to room temperature. The double helices
are generally limited to a few dozen (perhaps 50) glucose
residues, leaving great lengths of single helix that can entrap
enough iodine to give a strong blue color. Amylopectin retro-
gradation, in contrast, is more likely to involve chains from the
same molecule that are already much shorter than their amy-
lose counterparts. This process can take up to several days and
makes a translucent, soft paste. Loss of water from the gel or
paste during storage is termed syneresis.
Some of the components of a retrograded starch are so well
crystallized that they are resistant to enzyme attack and are
known as resistant starch (Table 3). These can contribute to
dietary fiber. Recrystallized autoclaved amylomaize starch is
sold as resistant starch by at least two manufacturers. Similar
CARBOHYDRATES | Starch: Chemistry 179
material is used as an additive to white bread by an Australian
bread manufacturer in order to increase insoluble dietary fiber
content without adversely affecting the color or texture of the
product.
Mutant Starches
Starch synthesis mutants have been well characterized in sev-
eral species. These have been valuable not only as industrial
and food components but also for elucidating the relative roles
of amylose and amylopectin in starch functionality.
GBSS-null mutants do not synthesize amylose (or do so to
only a limited extent), so the starch is 100% amylopectin and is
known as waxy on account of the appearance of the cereal
grains. These wx mutants have long been known in barley,
maize, and rice (glutinous) and have more recently been
developed in common wheat and pea. The gel temperature is
usually lower and peak viscosity higher in waxy starches than
in their normal counterparts. In many cases, final viscosity is
higher as well. The isolation of further waxy starches, for exam-
ple, from other grain legumes and pseudocereals, may give us
even more interesting properties.
High-amylose starches have been harder to develop, even
though one has been known for a very long time and was an
important part of Mendels original elucidation of the princi-
ples of genetics. The distinction between round and wrinkled
peas, called rugosus, is now known to be due to a mutation in
SBE I. The wrinkled phenotype is associated with an amylose
concentration of up to 80% in a reduced mass of starch. High-
amylose barley (amo-1) is 45% amylose, whereas high-amylose
maize starch can exceed 85% amylose. A barley containing no
amylopectin was developed by RNA interference with the SBEs.
Amylose appears to be digested more slowly than amylopectin
in the gut of monogastric animals. High-amylose starches have
been associated with increased satiety (i.e., reduced desire for
food), reduced glycemic index, and reduced insulin release in
comparison with normal-amylose starches. High-amylose
foods are thus an attractive option for people with diabetes
180 CARBOHYDRATES | Starch: Chemistry
and other metabolic problems related to insulin, glycemia, or
obesity.
Many mutations affect maize starch synthesis, including ae
(amylose extender), du (dull), h (horny), sh1 and sh2
(shrunken), and su1 and su2 (sugary). The double mutant ae
wx maize has no true amylose, owing to the wx mutation, but
has longer amylopectin chains (average 52 instead of 30 glu-
cose residues) than the wild type due to the ae mutation.
Starches of other double or triple mutants of maize have desir-
able properties that emulate or exceed those of chemically
modified starches. These include good freezethaw stability
(wx sh1) and low viscosity when hot during processing but
high viscosity when cool, which is desirable for canning (du
h, du ae, du su2, and ae su2). Starch of the ae du wx triple mutant
has high paste viscosity, shear resistance, and acid resistance,
and the paste has a creamy texture making it a suitable fat
substitute. Chemical analysis of chain length and branching
pattern has not yet been at sufficiently high resolution to
provide mechanistic explanations of these differences in pro-
cessing properties.
Conclusions
Numerous techniques have been used to elucidate aspects of
the physical and functional chemistry of starch. Although it is a
homopolymer consisting entirely of glucose, its structure is
extremely variable, showing differences in chain length,
branching pattern, and the incorporation of other molecules
such as proteins and lipids. These differences lead to a wide
range of processing properties. Aspects of the nomenclature
may be confusing to the novice, in particular the use of the
letters A, B, and C to indicate such diverse traits as different
types of amylopectin chain, granule size, or x-ray diffraction
pattern. Many questions remain to be answered about the
various levels of organization within the starch granule, includ-
ing the nature of the initiation of the starch granule itself.
Exercises for Revision
Distinguish between amylose and amylopectin.
What happens to starch during gelatinization and
retrogradation?
Describe the ABCs of starch granule size, chain length, and
x-ray diffraction.
Describe the various levels of crystallinity in a starch
granule.
Exercises for Readers to Explore the Topic Further
How are we manipulating the genes of starch synthesis to
make new types of starch for food and industrial purposes?
See articles on starch synthesis, uses of native starch, and
functional foods and various articles on breeding and
genetics of cereals, pulses, and pseudocereals.
What are the starch quality requirements (e.g., granule size
and amylose content) for bread, or biscuits, or noodles, or
pasta? See articles on the relevant food item.
The initiation of starch granules, particularly of the second-
ary starch granules in wheat, barley, and rye, remains an
enigma, as does the regulation of granule size and shape.
See also: Barley, Rice and Maize Processing: Barley: Malting;
Breeding of Grains: Barley: Genetics and Breeding; Maize: Breeding;
Wheat Breeding: Exploiting and Fixing Genetic Variation by Selection
and Evaluation; Carbohydrates: Carbohydrate Metabolism; Resistant
Starch and Health; Starch: Synthesis; Food Grains and Well-being:
Functional Foods: Overview; Genetics of Grains: Maize: Genetics;
Rice: Genetics; Processing of Grains: Starch: Modification; Starch:
Uses of Native Starch; The Cereal Grains: Durum Wheat: Overview;
Maize: Overview; Oats: Overview; Rice: Overview; Rye Grain: Its
Genetics, Production, and Utilization; Wheat: An Overview of the Grain
That Provides Our Daily Bread; The Legumes and
Pseudocereals: Amaranth: Overview; Beans: Overview; Chickpea:
Overview; Grain Legumes and Their Dietary Impact: Overview; Pea:
Overview; Pseudocereals: Overview; Quinoa: Overview; Wheat
Processing: Chemistry of Cake Manufacturing; Wet Milling of Wheat;
Wheat: Dry Milling; Wheat-Based Foods: Noodles: Starch; Pasta.
Further Reading
Atwell WA, Hood LF, Lineback DR, Varriano-Marston E, and Zobel HF (1988)
The terminology and methodology associated with basic starch phenomena.
Cereal Foods World 33: 306311.
BeMiller JN and Whistler RL (2009) Starch Chemistry and Technology, 3rd edn.,
904 pp. New York: Academic Press.
Bertolini AC (2009) Starches: Characterization, Properties and Applications, 288 pp.
Boca Raton, FL: CRC Press.
Donald AM, Perry PA, and Waigh TA (2001) The impact of internal granule structure on
processing and properties. In: Barsby TL, Donald AM, and Frazier PJ (eds.) Starch:
Advances in Structure and Function, pp. 4552. Cambridge: Royal Society of
Chemistry.
Morrison WR (1995) Starch lipids, starch granule structure and properties.
In: Schofield JD (ed.) Wheat Structure, Biochemistry and Functionality,
pp. 261270. Cambridge: Royal Society of Chemistry.
Parker R and Ring SG (2001) Aspects of the physical chemistry of starch. Journal of
Cereal Science 34: 117.
Sonnewald U and Kossmann J (2013) Starches From current models to genetic
engineering. Plant Biotechnology Journal 11: 223232.
Relevant Websites
http://www.corn.org Corn Refiners Association, Inc, Washington DC, USA.
http://www.jic.bbsrc.ac.uk The John Innes Centre, Norwich, UK.
http://www.mri.bund.de/en/de/institute/sicherheit-und-qualitaet-bei-getreide.html
Department of Safety and Quality of Cereals, Max-Rubner Institut, Detmold,
Germany.
http://www.starch.dk International Starch Institute, Aarhus, Denmark.
http://eu.wiley.com/WileyCDA/WileyTitle/productCd-2041.html Starch/die Starke
(Journal home page), Wiley Interscience Publishing.
 Starch: Synthesis
A Regina, CSIRO Food Futures National Research Flagship, Canberra, ACT, Australia; CSIRO Plant Industry, Canberra, ACT, Australia
S Rahman, Monash University Malaysia, Bandar Sunway, Malaysia
Z Li, CSIRO Food Futures National Research Flagship, Canberra, ACT, Australia; CSIRO Plant Industry, Canberra, ACT, Australia
MK Morell, International Rice Research Institute, Los Banos, Philippines
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Starch is the major component of grains.
It is important for human nutrition and also as raw material
for various domestic and industrial products.
Starch is a polymer of glucose with two different types of
linkages and other modifications.
Different plant starches can vary considerably in some of
their characteristics such as the size and shape of the
granule, ratio of different linkages, and proportion of
modifications.
Starch biosynthesis is carried out in special compartments
of the plant cell.
Many enzymes are involved in starch biosynthesis and the
roles of all of these enzymes have not yet been clarified.
Further knowledge of starch biosynthesis will enable the
production of starch more precisely tailored to specific
needs.
Learning Objectives
To appreciate the chemical nature of starch.
To appreciate the complex nature of the biosynthesis of
starch.
To appreciate gaps in knowledge and the potential for
targeting starch for specific end uses.
Introduction
Starch makes up by far the largest component of cereal grains
and is also found in fruits, roots, tubers, and leaves. It is a very
important source of energy in our diets. Most starch is easily
digested; however, some starches termed resistant starch are
digested only after they reach the large intestine, and recent
research shows that this type of starch has positive attributes
for bowel health. Starch also performs two basic roles in the
food industry: as a stabilizer providing viscosity, texture,
mouthfeel, and consistency to food products and as a proces-
sing aid to facilitate manufacturing. As a consequence of its low
cost and diverse functionality, starch is used in a range of food
and nonfood industries.
Starch is a polymer of glucose monomers linked by a-1,4
and a-1,6 linkages, so it is a glucan. Cellulose is also a glucan;
however, the single type of glucose linkage between the glucose
molecules (b-1,4) provides very different properties. The a and
the b refer to different spatial relationships between the glucose
residues. In starch, the frequency of a-1,6 linkages defines the
two major components of starch amylose and amylopectin.
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00094-2
Amylose and amylopectin differ in their degree of polymeriza-
tion (DP) or number of glucose residues and branching fre-
quency. Amylose is mainly linear with <1% a-1,6 linkages and
a DP of 5005000, whereas amylopectin is highly branched
with 46% a-1,6 linkages and a DP of 50 000500 000. The
amylose/amylopectin ratio and the details of the branching
(such as the length of the branches and distance between
branches) are important determinants of the suitability of
starch for specific end uses. Generally, the amylose content of
starch is
25%, but it can vary from 0% to 90% depending on
alterations in the starch biosynthetic pathway.
In nature, starch is found in granules. x-Ray diffraction
studies have revealed a crystalline structure for such granules,
with alternating crystalline and amorphous zones. The x-ray
diffraction pattern is determined by the proportion of the
external chains that are aligned in crystalline arrays and by
the packing in the array. Amylopectin molecules are radially
arranged with their nonreducing ends pointing toward the
surface. The location of amylose chains within this structure
is unclear, and it has been suggested that it is present in the
amorphous region and/or interspersed in the amylopectin
crystalline region.
Starch is synthesized in plants within organelles that are
either photosynthetic (such as the chloroplast of leaf cells) or
nonphotosynthetic (such as the amyloplast of cereal endo-
sperm cells). The starch in chloroplasts is synthesized during
the day and broken down at night, and it is often termed
transitory starch. In contrast, the starch synthesized in seeds
is broken down only on germination, and the starch is termed
reserve starch. There are some differences in the detail of bio-
synthesis between the two types of starch. This article is mainly
concerned with the starch deposition within the non-
photosynthetic amyloplast.
Starch from Different Cereals
Starch granules in the reserve tissues vary widely in size distri-
bution and shape between species (see Table 1 and Figure 1).
Maize and rice starches exhibit a unimodal granule size distri-
bution ranging from 5 to 20 mm for maize and 2 to 5 mm for
rice. Starches of wheat, barley, and rye are characterized by a
bimodal distribution composed of an A granule population of
1035 mm diameter, which are lenticular in shape with a char-
acteristic equatorial groove, and a B granule population of
generally spherical granules with a diameter less than 10 mm.
In barley, a trimodal distribution of starch granules with C
granules of diameters less than 5 mm is also reported. In wheat,
the larger granules, which are initiated during the early stages
of endosperm development, constitute 7080% of the starch
181
182 CARBOHYDRATES | Starch: Synthesis

Table 1 Typical properties of starch from different sources Waxy starches are also often preferred as thickeners because of
the greater solubility of starches without amylose. In cereals,
Starch Starch high-amylose starches are best known in maize among cereals
granule granule Apparent Gelatinization
where amylose content to 90% has been achieved. More
size range size amylose onset
recently, high-amylose wheat, barley, and rice have also been
Source (mm) distribution content temperaturea( C)
developed. The starch granules with elevated amylose content
Wheat 335 Bimodal 25 57 appear to be distorted. High-amylose starches are in demand
Barley 235 Trimodal 22 56 because such starches are useful starting points for the produc-
Rice 25 Normal 18 70 tion of resistant starches in foods and can be used in packaging
Maize 520 Normal 28 64 materials and adhesives.
a An important property of starch granules is the gelatiniza-
Gelatinization temperature measured by differential scanning calorimetry using starch
mixed with water in a ratio of 2 parts of water to 1 part of starch. The heating rate was
tion onset temperature, the temperature at which the granules
10  C min1 over a temperature range of 25150  C. begin to lose internal order and crystallinity. The gelatinization
Source: Data compiled from Jane J, Chen YY, Lee LF, et al. (1999) Effects of temperature of starches can vary considerably, from
57  C
amylopectin branch chain length and amylose content on the gelatinization and pasting for barley starches to over 75  C for rice. Mutations in some of
properties of starch. Cereal Chemistry 76: 629637; Rahman S, Li Z, Batey I, Cochrane the starch synthases (SSs) (GBSS and SSIIa) have been shown
MP, Appels R, and Morell M (2000) Genetic alteration of starch functionality in wheat. to lower the gelatinization onset temperature to
46  C in
Journal of Cereal Science 31: 91110. mutants of SSIIa in wheat and barley. On the other hand, the
suppression of starch branching enzyme (SBE) IIa and IIb
by weight, and the more numerous smaller granules, which are tends to increase gelatinization onset temperature to
60  C.
started later, contain only 2030% of the total starch in grain. The differences in gelatinization correlate with the differences
There have been some reports of slight variation in amylose in the branch lengths of starches from different sources and in
content between the A and B granules and between different the packaging of branches in the granule.
regions of the granule. Another important property is the viscosity of the gelati-
The amylose/amylopectin ratio does not normally vary nized starch. This is related to phosphate content and is a
greatly between sources, with a ratio of 1:3 being common. principal difference between starches from cereals and tubers.
However, mutations where amylose is completely missing are Potato starches have
0.5% phosphorylated glucose residues
fairly common and such starches are known as waxy starches. compared with
0.05% for cereal starches. Usually, the
In some food preparations, such starches are preferred, for phosphate is linked to the carbon at position six in glucose
example, in the preparation of many Asian sticky rice desserts. molecules. The difference in the phosphorylation imparts

Wheat Barley

Maize Rice
Figure 1 Starch granules from common cereals. Scanning electron micrographs by CSIRO Plant Industry by scanning electron microscopy.

tuber starches with properties significantly different to cereal
starches in terms of viscosity and flow properties. It has also
been shown that starches with more short and intermediate
glucan chains have a high starch viscosity.
The lipid content of cereal starches is low,
1%. However,
even this can be divided into three operational classes: non-
starch, surface, and internal lipids. The internal lipids of wheat
consist entirely of lysophospholipids, which are complexed
inside linear glucan chains. The fatty acid composition of lipids
differs slightly between wheat and rice, with wheat being richer
in 18:2 (linoleic acid) and rice having more 16:0 (palmitic
acid) fatty acids. How lipids accumulate in cereal starches is
not yet clear, but the hydrophobic environment in the interior
of the glucan chain leads to complexes of lipids with amylose.
Pathway from Sucrose to Starch Granule
Pathway Preceding Starch Biosynthesis
Starch synthesis requires a hexose phosphate supply. Plants
produce sugars in photosynthetic tissues (e.g., leaves and peri-
carp of the grain) where they can support starch synthesis. Some
of these sugars are exported as sucrose. In the endosperm,
incoming sucrose is either converted into uridine diphosphate
(UDP)glucose and fructose by a UDP-dependent sucrose
synthase or converted to hexoses by invertase, as observed in
maize. The UDPglucose or hexose phosphate derived from
fructose is interconverted to glucose-1-phosphate by other
enzymes such as UDPglucose pyrophosphorylase, hexoki-
nase, and phosphoglucomutase. This glucose-1-phosphate
can be used in various metabolic pathways; however, once the
glucose-1-phosphate is utilized to form adenosine diphosphate
(ADP) glucose (ADPG), a commitment is made to the synthesis
of starch.
Biochemistry of Starch Biosynthesis
A consensus view of starch biosynthesis in cereal endosperm is
shown in Figure 2. Starch biosynthesis requires four types of
enzymes, which are as follows:
Isoamylase
SBE IIb
SBE IIa
ADPG SBE I
75% Amylopectin
PPase
SS III
ATP PPi SS II
SS I
G1P ADPG 25% Amylose
ADP
ADPG GBSS
Amyloplast
Figure 2 A consensus view of the starch biosynthetic pathway in
cereals.
CARBOHYDRATES | Starch: Synthesis 183
(1) ADPG pyrophosphorylase, which produces ADP-glucose
from glucose-1-phosphate and ATP.
(2) SSs, which elongate glucan chains by using ADP-glucose to
add a glucose residue at the nonreducing end.
(3) SBEs, which introduce a-1,6 branches to the glucan chain.
These branches can then be extended by SSs.
(4) Debranching enzymes remove branches from starch mol-
ecules. The role of debranching enzymes in starch synthe-
sis is still being debated, although the genetic evidence for
a role for these enzymes is unambiguous.
Other enzymes are also capable of influencing the properties of
the starch produced and these will be discussed later.
Maize has been the most extensively characterized model
system for the isolation of mutations affecting starch biosyn-
thesis. Analysis of these mutants (along with those from other
model systems such as Arabidopsis and Chlamydomonas) has
greatly contributed to the exploration of the key enzymes and
their isoforms involved in starch biosynthesis. In maize, muta-
tions in genes affecting starch synthesis such as the waxy and
amylose extender mutations can be brought about by the
insertion of transposable elements, a DNA sequence capable
of moving from one location to another within a genome.
Such mutations often lead to alterations in the structure and
properties of starch. Specialty maize varieties such as waxy
(granule-bound starch synthase (GBSS) I mutant), amylose
extender (SBEIIb mutant), sugary-2 (SSIIa mutant), sweet
corn (sugary-1, isoamylase mutant), and dull (SSIII mutant)
find applications for specific food and industrial uses.
ADPG Pyrophosphorylase
ADPG pyrophosphorylase is believed to be the key enzyme
that determines the rate of starch synthesis in leaves. Genetic
engineering has been used to introduce the ADPG pyropho-
sphorylase from Escherichia coli (which is not subjected to
feedback inhibition) into potatoes, and tubers with a higher
content of starch were obtained. It is not clear, however,
whether ADPG pyrophosphorylase is also the rate-limiting
enzyme for starch biosynthesis in the grain, as the purified
enzyme from leaves and grains shows differences in properties.
ADPG pyrophosphorylase from both leaves and endosperm
is composed of four subunits, in which two of each type are
large and small. The large subunit has a mass of
55 kDa,
whereas the small subunit is slightly smaller,
50 kDa. The
sequence of amino acids of these subunits is
50% identical.
Mutations in the sequence of either of these subunits can
destroy or reduce the activity of the enzyme, and then grains
or seeds with a shrunken appearance (due to decrease in starch
synthesis) are produced. Such mutations in ADPG pyropho-
sphorylase were given the names brittle-2 (mutation in small
subunit) and shrunken-2 (mutation in large subunit) when
they were discovered in maize (Table 1).
Recent evidence shows that ADPG pyrophosphorylase in
the cereal grain exists in both the cytoplasm and amyloplast of
endosperm cells; however, most of it is in the cytoplasm. This
enzyme in the cereal grain differs from that in the seeds of
dicots and leaves of all the plants investigated so far, where
ADPG pyrophosphorylase appears to exist exclusively in the
184 CARBOHYDRATES | Starch: Synthesis

plastids. Evidence from both monocots (such as barley and
maize) and dicots (such as peas) suggests that there is more
than one gene for the large and small subunits of ADPG
pyrophosphorylase, and different versions of these may code for
the forms that are expressed in the cytoplasm and in the plastid.
The annotation of the rice genome sequence also indicates
multiple gene sequences coding for ADPG pyrophosphorylase.
The transfer of ADP-glucose from the cytoplasm of endo-
sperm cells to the amyloplast is important as the other enzymes
in the process are located there. The Brittle-1-encoded ADPG/
ADP antiporter facilitates this transfer in maize. Mutations are
known where this transfer is affected and this too produces
grain where starch synthesis is affected (Table 2).
ADPG pyrophosphorylases exhibit complex regulatory
properties that vary depending on the species and tissue
source. The enzymes from the leaves of all plants examined,
maize endosperm, and potato tubers are allosterically activated
by 3-phosphoglyceric acid and inhibited by inorganic phos-
phate. Pea and bean embryo and wheat and barley endosperm
ADPG pyrophosphorylases, on the other hand, are much less
sensitive, or insensitive, to modulators compared with their
leaf counterparts. The allosteric properties of ADPG pyropho-
sphorylases are of potential importance in determining its role
in controlling the rate of starch synthesis. Altered starch bio-
synthetic rates resulting from altered allosteric properties due
to mutations in ADPG pyrophosphorylase subunits have been
demonstrated in maize, barley, and the green algae Chlamydo-
monas. Alterations in the properties of ADPG pyrophosphory-
lase have been utilized in the plants grown in the greenhouse
to increase starch production and thus seed weight.

Table 2 List of selected starch mutants

Plant Causal mutations/only enzyme

Mutant Phenotype species affected

Brittle-1 (bt1) Low starch Maize Adenylate transporter

Shrunken-1 Low starch Maize Sucrose synthase
(sh1)
Rugosus 4 Low starch Pea Sucrose synthase
(rug4)
Rugosus 3 No starch Pea Phosphoglucomutase
(rug3)
Rb Decreased starch content and increased levels of sucrose Pea ADPG pyrophosphorylase
Brittle-2 (bt2) Low starch Maize Small subunit of ADPG
pyrophosphorylase
Shrunken-2 High levels of sucrose and low levels of starch Maize Large subunit of ADPG
(sh2) pyrophosphorylase
Waxy (wx) Zero amylose Maize, GBSSI
wheat
Low amylose Low level of amylose Pea GBSS
(lam)
Shrunken (shx) Reduced starch content, altered granule size distribution Barley SSI (not known to be casual)
Sugary-2 (su2) Altered granules, high sugar Maize Starch synthase IIa (?)
Rugosus 5 Granules with deeply divided lobes, reduced amylopectin synthesis, altered Pea SSII
(rug5) amylopectin chain length
Dull (du1) Mature kernels with tarnished, glassy, and dull appearance. High apparent Maize Mutation at SSIII, SBEIIa affected
amylose content secondarily
Amylose High levels of amylose Maize, SBEIIb, RBEIII
extender (ae) rice
High amylose Higher level of amylose Barley Unknown
(amo1)
Floury-2 (flo-2) Soft, white endosperm that crumbles easily into powder. High amylose content Rice RBEI, RBEIII, and GBSS
(not known to be casual)
SbeIIa::mu No endosperm phenotype, altered leaf starch Maize BEIIa
Sbe1::mu No known phenotype Maize BEI
Rugosus (r) Wrinkled seed, lowered starch, and enhanced amylose, sucrose, and lipid levels Pea SBEI
Sugary-1 (su1) Low starch, phytoglycogen accumulation Maize, Isoamylase
rice
Soft starch (h) Loosely packed, large granules Maize Unknown
SGP-1 Abnormal starch granule morphology. High amylose content Wheat SSII
Isoamylase Phytoglycogen accumulation. No B granule initiation Barley Isoamylase
mutant
Sex6 Decreased amylopectin synthesis, shortened amylopectin chain length Barley SSIIa
distribution, reduced gelatinization temperature

Source: Data compiled from the references in Further Reading section. ? not clear if the gene responsible has been identified.

Starch Synthases
SSs elongate preexisting glucan chains in the amyloplast by
adding the glucosyl moiety from ADP-glucose to the nonredu-
cing end of an existing a-1,4-glucan.
Ten different SS genes have been identified in plants that
encode two classes of SSs. One class of SS is only found bound
to starch granules and these are known as GBSS. The other class
of SS is present either in the amyloplast stroma alone or dis-
tributed between the stromal and granular fractions, and these
are known as soluble SSs. Four types of soluble SS have been
reported in crop plants: SSI, SSII, SSIII, and SSIV. In monocots,
the SSII has been further subdivided into the IIa and IIb forms,
SSIII into SSIIIa and SSIIIb forms, and SSIV into SSIVa and
SSIVb forms. The primary structures of soluble SSs from different
sources share a distinguishing feature, an N-terminal extension
with little homology between sequences from different sources.
Comparison of the deduced amino acid sequences of SSs
revealed that maize SSI, SSIIa, SSIIb, and SS IIIa contained
N-terminal extensions of 93, 176, 144, and 656 amino acids,
respectively, compared with GBSS. Pea SSII also contains a flex-
ible 162-amino acid N-terminal arm. It has been shown through
expression analysis using N-terminally truncated SSII that the
N-terminal extension is not essential for the catalytic activity of
the enzyme but is probably related to substrate binding.
There are two types of GBSSs: I and II. GBSSII is similar in
amino acid sequence to GBSSI but is not expressed in the grain.
GBSSI is a single polypeptide,
60 kDa in mass. If the grain
lacks an active GBSSI, then starch is composed almost entirely
of amylopectin. Such starch is called waxy starch (Table 2).
Clearly then, GBSSI is essential for the synthesis of amylose
and the other SSs cannot substitute for it.
The hexaploid nature of wheat allows a range of starches of
intermediate amylose content to be developed, an opportunity
not possible in diploids such as barley. In wheat, the GBSSI
gene has been mapped as a triplicate set of single-copy home-
oloci on chromosome arms 7AS, 4AS, and 7DS. A reduction in
the proportion of amylose is observed when the GBSSI activity
is reduced, and these starches are known as low-amylose or
partial waxy starches. The loss of only one of the three iso-
forms, the 4A form of GBSS, has clearly been shown to yield a
starch with increased ability to swell in water on heating. This
property is desirable for udon noodle production, and such
low-amylose starches from wheat are preferred for this end use.
The soluble SSs are also all single polypeptide enzymes. SSI
is
75 kDa in mass and is found both free in the amyloplast
and bound to the starch granule. Starches of SSI mutants of
Arabidopsis and rice and SSI RNAi silenced wheat have reduced
amounts of amylopectin, low peak viscosities, and high peak
gelatinization temperatures. Studies in these modified lines
have shown that SSI is involved in the synthesis of short chains
of the amylopectin (DP 812).
SSII has been subdivided into IIa, IIb, and IIc forms on the
basis of gene sequence comparisons; however, the IIb and IIc
forms appear not to be expressed or expressed at very low
concentrations in the cereal endosperm. The IIa form is

85 kDa in mass, although the wheat enzyme appears to be
over 100 kDa by electrophoresis. Barley and wheat lines lack-
ing SSIIa have been produced, and the starch from these lines is
CARBOHYDRATES | Starch: Synthesis 185
clearly different from normal. The barley mutants have high
amylose (70%); the wheat mutants, less so (35%). It is not
clear if there are differences in the structure of the amylose. The
starch granules are of distorted appearance and have a lower
gelatinization temperature. The proportion of branches of
725 glucose units in length is decreased, while the proportion
of branches of 46 glucose units in length is increased. There
are also effects on the abundance of other starch biosynthetic
enzymes, so it is difficult to know how many of the effects on
starch properties are the direct result of the lack of this enzyme
and how many are due to the other enzymes affected. The
differences in the starch between indica and japonica rice
cultivars appear to be due to a combination of alterations in
SSIIa and GBSS.
SSIII has been subgrouped into SSIIIa and SSIIIb. SSIIIb is
mainly expressed in the pericarp and does not appear to be
easily detected in the endosperm. SSIIIa is a polypeptide of

180 kDa. The absence of this enzyme in maize leads to a
slightly higher amylose content, and the starch looks dull
compared with normal. Mutants in rice and barley have a
reduced amylopectin content, a reduced starch content, but
an increased amylose content. The mutation of SSIIIa increases
the expression of SSI and GBSSI in the endosperm. An inter-
mediate glucan fraction (
15%), which is distinguished from
amylose and amylopectin, occurs in starch from maize lines
missing SSIIIa.
SSIV has been subdivided into SSIVa and SSIVb isoforms in
cereals. Both isoforms are expressed in the endosperm,
although SSIVa is expressed at a higher concentration than
SSIVb in rice. The importance of either of the SSIVs for starch
synthesis in the endosperm is unknown at present. Mutation of
SSIV in Arabidopsis dramatically reduces the number of starch
granules per plastid, leading to a large increase in starch gran-
ule size in the leaf, and it appears that SSIV has a critical role in
regulating the number of starch granules.
Starch Branching Enzymes
SBEs, as their name suggests, are required for the addition of
branches to linear glucans. Their impact is clearly more on the
synthesis of amylopectin than amylose, although branching
enzymes may also have a subtle role in amylose synthesis.
There are two broad types of SBEs: SBEI and SBEII. Expres-
sion of these enzymes in bacteria has shown that SBEI adds
longer branches than SBEII. They are both single polypeptide
chains of
85 kDa. SBEI and SBEII share
60% sequence
identity over the middle third of the molecules.
The importance of SBEI for starch biosynthesis in the grain
is not clear. Mutations in SBEI in maize and wheat did not
produce a clear phenotype. A number of isoforms of SBEI may
exist in wheat. However, in rice, an SBEI mutant contains
amylopectin with significantly decreased proportions of long
chains of DP 37 and above and short chains of DP 1221.
However, the proportion of short chains of DP 10 and below
and intermediate chains of DP 2434 increased. Starches from
the rice SBEI mutant had a low gelatinization onset tempera-
ture. SBEI in rice may also play a crucial role in the synthesis of
186 CARBOHYDRATES | Starch: Synthesis
amylopectin molecules by interacting with phosphorylase 1 to
form branched maltodextrin primers.
SBEII is divided into two isoforms in cereals: IIa and IIb.
The amino acid sequences of IIa and IIb isoforms are very
similar (over 80% sequence identity within a species). In
maize, the IIb form makes up
90% of the starch branching
enzyme II activity in the endosperm, but the proportion of the
two isoforms is nearly equal in wheat. Maize lines lacking
starch branching enzyme IIb produce a very high (70% and
higher) proportion of amylose. Rice SBEIIb mutants also con-
tain high-amylose starch, but the proportion is much lower
(
40%) than in maize. In wheat and barley, suppression of
SBEIIb expression using RNAi technology has no effects on
starch amylose content. Suppression of both SBEIIa and SBEIIb
is required for the production of amylose (>70%). In rice
suppression of SBEI, IIa and IIb are required to achieve amy-
lose contents of more than 50%. High-amylose starches are
linked to the formation of resistant starches in products. Resis-
tant starch has been shown to have very beneficial effects if
present in the diet.
Debranching Enzymes
Debranching enzymes remove branches from branched glu-
cans. Two types of debranching activities, pullulanase and
isoamylase, have been described from developing endosperms
of rice and maize. They differ in the type of glucan they can
debranch with pullulanase debranching more efficiently
open structures (e.g., pullulan) and isoamylase debranching
the more branched structures (e.g., glycogen). The role and
importance of these enzymes for starch biosynthesis are still
being debated.
Pullulanase is a single polypeptide enzyme of
100 kDa in
mass. Grains from a rice pullulanase mutant contain starch
with a reduced proportion of short chains (DP 13 and
below) and a slightly lower gelatinization temperature (lower
by 13  C). Such mutations in rice do not change the amylose
content. A pullulanase mutant in maize contains significant
quantities of low-molecular-mass a-1,6-polyglucans in the
endosperm. Similar to rice, mutants in maize pullulanase con-
tain starch with unaltered amylopectin structures and amylose/
amylopectin ratios.
Isoamylase activity in plants is controlled by three gene
products, designated as ISA1, ISA2, and ISA3. Of these, ISA1
and ISA2 are involved in starch biosynthesis and ISA3 is
involved in starch breakdown. ISA1 is more catalytically active
than ISA2 and can form complexes with it. ISA1 is a single
polypeptide enzyme of
90 kDa in mass. It can debranch
highly branched structures like amylopectin and glycogen.
The lack of this enzyme in rice and maize is associated with
the production of a very highly branched and unusual poly-
saccharide called phytoglycogen. Such lines also contain high
levels of free sugars and are known as sugary mutants. How-
ever, there is also a reduction in the level of pullulanase in
these sugary mutants, so it is difficult to know which of the
effects are directly due to the lack of ISA1. The effect of the loss
of ISA2 is slight in rice.
There are differences between the isoamylase mutants
described in rice, maize, and barley. In rice, different phenotypes
were observed for the sugary mutants ranging from no starch
and only phytoglycogen to starch in the outer layer of the
endosperm and phytoglycogen internally. In contrast, in barley,
both phytoglycogen and starch were reported in the same
endosperm cells. The maize sugary mutants also contain both
phytoglycogen and starch. In barley, it has been reported that
the lack of isoamylase leads to an alteration in the initiation
pattern of starch granules.
While it is established that isoamylase plays a significant
role in starch biosynthesis, the mechanism by which it acts is
not resolved. Two mechanisms have been proposed. One pro-
posal known as the glucan trimming mechanism suggests that
isoamylase removes moderately and loosely spaced branches
from a soluble preamylopectin structure that interferes with
the crystallization of amylopectin and promotes the formation
of a stable amylopectin structure with regularly packed glucose
units. In the absence of isoamylase activity, the preamylopectin
structure is further branched by branching enzymes and results
in the production of phytoglycogen. The second model is
based on the assumption that a competition exists between
polysaccharide aggregation into starch and the nonproductive
formation of water-soluble polysaccharides (WSPs), in turn
resulting in a competition for both carbon source and soluble
enzymes involved in amylopectin synthesis. The role of isoa-
mylase here would be to clear the stroma of WSPs and prevent
phytoglycogen formation. A role in starch granule initiation
and growth, apart from phytoglycogen suppression, for isoa-
mylase is also suggested based on the analysis of barley
isoamylase mutants.
Other Enzymes with Potential Roles in Synthesis
Many of the new insights into starch synthesis have come from
the study of simpler systems, such as Chlamydomonas and
Arabidopsis. Such studies have indicated other loci that are
important in starch synthesis in these organisms. It is possible
(although by no means certain) that genes encoded by such
loci are also important for starch synthesis in the grains of
cereals.
D-Enzyme or disproportionating enzyme has the property
of breaking a glucan chain and adding one portion to the
nonreducing end of a preexisting chain. In this way, one
chain is made shorter but the other one is made longer. In
Chlamydomonas mutants where the gene for D-enzyme is
affected, the starch produced appears to have somewhat more
amylose. These mutants are required to be grown in constant
light. Genes for D-enzyme have been described from Arabidop-
sis, rice, maize, barley, and wheat.
Starch phosphorylase is another enzyme that could have an
impact on starch biosynthesis in the grain. This enzyme breaks
down starch and was originally considered to be important for
degradation. There are at least two isoforms in the plant a
cytosolic form and a plastidic form; however, Chlamydomonas
appears to contain three isoforms. The lack of one of these
isoforms leads to a starch excess phenotype and the production
of larger starch granules. In rice, the same phenotype is
observed, but this is more obvious at low growth temperatures
(
20  C). In wheat, plastidic phosphorylase appears to be

associated with leaf and seed endosperm accumulation, not
with the degradation of reserve starch.
The initiator protein for glycogen is called glycogenin. Glyco-
genin is an autocatalytic glycoprotein, and the glucose residues
provide the starting point for glycogen synthesis. By analogy,
with this, a number of glycogenin-like proteins have been
described from plants, but again, the importance of these pro-
teins to starch biosynthesis in the grain is yet to be elucidated.
The R1 protein is important for starch phosphorylation in
potatoes. If this protein is downregulated in potatoes, then the
starch produced is sharply reduced in phosphate content and
increased in amylose content. The peak viscosity was also
reduced. As cereal starch contains naturally far less phosphate,
the importance of this protein for cereal starch biosynthesis
needs to be ascertained. However, sequences similar to the
gene for the R1 protein are found in the cereal genome. In
Arabidopsis, potato, and wheat, the absence of the R1 protein
leads to a decrease in starch breakdown and the consequent
production of the starch excess phenotype.
Protein Interactions in Starch Biosynthesis
Recent studies in maize and wheat have shown that the key
enzymes involved in starch biosynthesis congregate into protein
complexes during starch biosynthesis. These complexes may act
as functional units to perform the task of granule formation.
Several such interactions are now known. For example, SSIIa
forms a complex with SSI and SBEIIb in maize. The composition
of such assemblies may change when there are changes in the
expression of key enzymes. For example, in the amylose
extender mutant of maize, in which the SBEIIb activity is lack-
ing, SBEI, SBEIIa, and starch phosphorylase are recruited into
the complex involving SSIIa and SSI.
Genes and Genetics in Wheat
The structure of the genes for the principal starch biosynthetic
enzymes mentioned earlier has been described from wheat
(Figure 3). The genes are all complex in intron/exon structure.
The genes differ considerably in length, varying from 6 to 11 kb.
GBSS
SSI
SSII
SSIII
SBEI
SBEIIa
SBEIIb
Isoamylase
0 2 4 6 8 10 11 12
Figure 3 Structure of wheat starch biosynthetic genes (data from
authors laboratory). Exon sequences are illustrated by the thick lines;
intron sequences by the narrow line.
CARBOHYDRATES | Starch: Synthesis 187
The processing of the initial transcript to the final RNA is
clearly a complex process. Many of the mutants in GBSSI are
due to errors or inefficiencies in the processing of the RNA. In
barley, the waxy mutations have been reported that produce
low amylose, rather than zero amylose. In these mutant starch
granules, the outer cell layers of the endosperm contain more
amylose than the inner layers. Investigation of these mutants
revealed that the change in expression of GBSSI in these lines is
due to a 413-bp deletion of part of the promoter and 50 -
untranslated region of the gene. Two other barley waxy families
analyzed have no detectable level of amylose due to a 1-bp
alteration in the GBSS gene that completely eliminates GBSS
activity. In rice, a single nucleotide polymorphism at the leader
intron 50 splice site of the GBSS gene has been shown to be
associated with changes in amylose content due to altered
efficiency of GBSS mRNA processing. A sequence of AGGTATA
at this site is found to be associated with cultivars having
an amylose content of above 18% as against a sequence of
AGTTATA that is present in cultivars having lower amylose
content.
The locations of most of the starch biosynthetic enzyme
genes are known in wheat and shown in Figure 4. Many of
these genes are on chromosome 7. In rice, the locations of the
gene are known from the genome sequence, and the locations
of the wheat genes are consistent with the established relation-
ships of the chromosomes in these two grasses.
Future Work
Genomics has essentially provided the whole genome
sequence of Arabidopsis and rice. The sequencing of the wheat
and barley genomes is also nearing completion. It should soon
be possible to identify all putative starch biosynthetic enzyme
genes in the genomes of these species.
The function of such putative candidate genes can be ana-
lyzed by performing conditional knockouts of selected genes,
and powerful RNAi technology can be used. TILLING has also
been used to obtain desired mutants in starch biosynthesis.
Additionally, more sophisticated gene-editing techniques
using DNA nucleases based on technologies such as TALENs
and CRISPRs may be applicable for such experiments. Con-
versely, gene inactivation through tagging will also yield infor-
mation about the role of candidate genes. This can arise either
through the identification of the tagged gene and analysis of
the phenotype or by observing a phenotype and then identify-
ing the gene inactivated. Both approaches will be highly
informative.
Analysis of expression of all of the genes in a tissue by
means of microarrays and transcriptomics can also be per-
formed. The ability to study the pattern of transcription of
the whole genome by means of chips and the transcriptome
will facilitate the study of linked genes and pathways. If tagged
mutants are compared with the wild type, then the total
changes in transcription can be identified, and pathways can
begin to be deciphered. For untagged mutants or natural phe-
notypes, all the genes affected in a particular phenotype can be
identified from microarray or sequence abundance analysis of
the expressed genes in bulked segregants differing in the phe-
notype studied.
188 CARBOHYDRATES | Starch: Synthesis

WxB1

WxA1 WxD1

SSI
SSII
Isoamylase
SSIII SBEIIb
SBEIIa AGP-S

SBEI

1 2 3 4 5 6 7
Figure 4 Chromosomal location of wheat starch biosynthetic genes. Data from authors laboratory observations and from the suggested readings.

New Starches in Cereals
It is clear that with increasing knowledge of the enzymes
involved in starch biosynthesis, the ability to alter the starch
produced is also gained. New products and processes will need
to be developed to capture any advantages occasioned by these
new starches. Although such new phenotypes can be produced
most easily by using transgenic technology, current public
opinion would make it prudent to also produce starches by
conventional breeding. The development of molecular
markers and rapid breeding technologies (such as the produc-
tion of doubled haploids and embryo culture and the ability to
quickly extract DNA from small samples) will undoubtedly
also speed up the production of novel types of starch by
conventional breeding strategies.
able to precisely manipulate the starch structure and function-
ality to suit specific end uses.
Exercises for Revision
Why is starch important for our daily life?
How many classes of enzymes are involved in starch
synthesis?
Which starch synthetic enzymes are important in the deter-
mination of starch structures?
Which starch synthetic enzymes are important in the deter-
mination of starch properties?
How can novel starches be produced in cereals?
What approaches can be used for the study of the roles of
the enzymes in starch synthesis?

Conclusion Exercises for Readers to Explore the Topic Further

The involvement of four classes of enzymes ADPG pyropho-
sphorylase, SSs, SBEs, and debranching enzymes in starch
biosynthesis is clearly established, although the precise roles of
isoforms in many cases are not clear. Other enzymes may also
have roles in starch biosynthesis. Starch phosphorylase and
disproportionating enzyme (D-enzyme) are examples. The
role in starch degradation of both of these enzymes is well
known. However, their involvement in starch synthesis,
although demonstrated from a few studies, is a matter requir-
ing further research. The priming of polysaccharide synthesis
and granule formation during starch biosynthesis is still
unresolved.
Although it is evident that mutations in the core genes lead
to specific starch phenotypes over a variety of species, species-
specific characteristics are observed that are often significant
and of practical importance. Understanding the complex
nature of starch biosynthesis has always been facilitated and
accelerated through investigations on model systems such as
Chlamydomonas and Arabidopsis and increasingly rice. However,
attempts should also be focused on individual species to be
The roles of different isoforms of starch synthetic enzymes
in the determination of starch structures and properties.
Interaction of the isoforms of starch synthetic enzymes in
starch synthesis.
Regulation of the isoforms of starch synthetic enzymes in
starch synthesis.
See also: Carbohydrates: Cereals: Chemistry and Physicochemistry
of Non-starchy Polysaccharides; Grains other than Cereals: Non-starch
Polysaccharides; Starch: Analysis of Quality; Starch: Chemistry;
Processing of Grains: Starch: Uses of Native Starch; Wheat-Based
Foods: Noodles: Starch.
Further Reading
Buleon A, Gallant DJ, Bouchet B, et al. (1997) Starches from A to C: Chlamydomonas
reinhardtii a model system to investigate the biosynthesis of the plant amylopectin
structure. Plant Physiology 115: 949957.

Gilbert RG, Wu AC, Sullivan MA, Sumarriva GE, Ersch N, and Hasjim J (2013)
Improving human health through understanding the complex structure of glucose
polymers. Analytical and Bioanalytical Chemistry 405: 89698980.
James MG, Denyer K, and Myers AM (2003) Starch synthesis in the cereal endosperm.
Current Opinion in Plant Biology 6: 215222.
Jane J, Chen YY, Lee LF, et al. (1999) Effects of amylopectin branch chain length and
amylose content on the gelatinization and pasting properties of starch. Cereal
Chemistry 76: 629637.
Morrison WR (1988) Lipids in cereal starches: A review. Journal of Cereal Science 8: 115.
Morrison WR (1993) Seed storage compounds: Biosynthesis, interactions and
manipulation. 12. Cereal starch granule development and composition.
In: Shewry PR and Stobart K (eds.) Seed Storage Compounds-Biosynthesis,
Interaction and Manipulation. Proceedings of the Phytochemical Society of Europe,
pp. 175190. Oxford: Oxford University Press.
Nakamura Y (2002) Towards a better understanding of the metabolic system for
amylopectin biosynthesis in plants: Rice endosperm as a model tissue. Plant Cell
Physiology 43: 718725.
Shewry PR and Morell M (2001) Manipulating cereal endosperm structure,
development and composition to improve end-use properties. In: Shewry PR,
Lazzeri PA, and Edwards KJ (eds.) Advances in Botanical Research Incorporating
Advances in Plant Pathology Biotechnology of Cereals, vol. 34, pp. 165236.
San Diego: Academic Press.
Tetlow IJ (2011) Starch biosynthesis in developing seeds. Seed Science Research
21: 532.
Thitisaksakul M, Jimenez RC, Arias MC, and Beckles DM (2012) Effects of
environmental factors on cereal starch biosynthesis and composition. Journal of
Cereal Science 56: 6780.
CARBOHYDRATES | Starch: Synthesis 189
Topping DL and Clifton PM (2001) Short-chain fatty acids and human colonic function:
Roles of resistant starch and nonstarch polysaccharides. Physiological Reviews
3: 10311064.
Tuncel A and Okita TW (2013) Improving starch yield in cereals by over-expression of
ADPglucose pyrophosphorylase: Expectations and unanticipated outcomes. Plant
Science 211: 5260.
Utsumi Y, Utsumi C, Sawada T, Furjita N, and Nakamura Y (2011) Functional diversity
of isoamylase oligomers: The ISA1 homo-oligomer is essential for amylopectin
biosynthesis in rice endosperm. Plant Physiology 156: 6177.
Yu TS, Kofler H, Hausler RE, et al. (2001) The arabidopsis sex 1 mutant is defective in
the R1 protein, a general regulator of starch degradation in plants, and not in the
chloroplast hexose transporter. Plant Cell 13: 19071918.
Zeeman SC, Kossmann J, and Smith AM (2011) Starch: Its metabolism, evolution, and
biotechnological modification in plants. Annual Review of Plant Biology
60: 209234.
Relevant Websites
http://www.ncbi.nml.nih.gov A resource for molecular biology information.
http://www.tigr.org/tdb Provides databases containing DNA and protein sequences
and gene expression, protein family and taxonomic data for microbes, plants and
animals.
 Starch: Starch Architecture and Structure
V Vamadevan, University of Minnesota, St. Paul, MN, USA; Agriculture and Agri-Food Canada, Guelph, ON, Canada
Q Liu, Agriculture and Agri-Food Canada, Guelph, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Granule morphology, structure, and composition of
starches from different genetic backgrounds.
Organization of glucan chains at different structural levels
of starch granules.
Cluster model of amylopectin versus building block back-
bone model.
Unit chain categories of amylopectin.
Branching point distribution pattern in amylopectin.
Structural parameters of amylopectin.
Learning Objectives
To understand the complex hierarchical structure of starch
granules.
To understand the concepts underlying proposed amylo-
pectin models.
To understand the internal structure of amylopectin.
To identify unresolved structural aspects of starch granules.
Introduction
Produced by plants as an end product of the photosynthetic
process, starch is one of the most abundant renewable bio-
polymers on Earth. It serves as a major energy source in the
human diet and an important functional ingredient in many
food and nonfood products. Principal sources are cereal grains
(e.g., wheat, maize, rice, and barley), tubers (e.g., potato),
roots (e.g., cassava), and pulses (e.g., pea, chickpea, and
bean). Starch is stored in the form of granules whose morphol-
ogy, structure, and composition are dependent on the genetic
background of the source.
Starch is synthesized in organelles called amyloplasts, and
the so-called hilum is thought to be the initiation point of the
biosynthesis of the starch granule. The position of the hilum in
the granule varies according to the botanical source. Starch
granules are classified as simple, compound, or semi-
compound types depending on how many granules are initi-
ated in each amyloplast. In simple starch granules (e.g., maize,
wheat, and barley), one granule is initiated in each amyloplast,
whereas compound granules (e.g., rice and oat) are composed
of multiple granules, known as subgranules, that are initiated
simultaneously in a single amyloplast. Semicompound gran-
ules (e.g., wrinkled pea) have two or more fused subgranules.
The molecular mechanism underlying starch granule initiation
is still largely unknown.
Starch Granule Morphology
The diameter of starch granules varies from 0.1 to 200 mm.
However, granules in the range of 2100 mm are the most
190 Encyclopedia of Food Grains, Second Edition
common. Cereal starches are generally small and have round
or polyhydric shapes with a concentric hilum and a unimodal-
size distribution. Wheat, triticale, barley, and rye starches,
however, exhibit bimodal size distributions, with large
(A-type) and small (B-type) granules. Starch granules from
tubers are usually large and oval-shaped with an eccentric
hilum. Pulse starches are normally medium-sized, with a bean-
like or ovoid shape and an elongated or starred hilum. High-
amylose starches generally appear as irregular, filamentous, or
rod-shaped granules (Figure 1).
Corn, sorghum, millet, wheat, rye, and barley starch gran-
ules are characterized by nonuniformly distributed surface
pores. Some granules have numerous pores, others feature
only a few, and the starch granules of rice, oat, pulse, potato,
tapioca, canna, yam, and arrowroot appear to have none at all.
In some starch granules, such as maize, sorghum, and millet,
radially oriented channels that connect to a central cavity have
been observed. Which biomechanism determines the size of
the granules in different plants and how the pores and chan-
nels are formed are still unclear.
Structural Components
Starch is synthesized as semicrystalline granules made up
of two principal types of glucan polymers: amylose and amy-
lopectin. Both are composed of anhydrous glucopyranose
units connected through a-(1,4)- and a-(1,6)-glycosidic
linkages. A third type of glucan polymer, known as an inter-
mediate material, is found in amylose extender (ae) maize
mutant starches and in some regular starches and starches
derived from developing or immature grains. Most starches
consist of 2030% amylose and 7080% amylopectin, with a
very low amount of intermediate material content. However,
amylose-free waxy starches contain essentially 100% amylo-
pectin, and high-amylose starches consist of < 50%
amylopectin.
The frequency of the a-(1,6)-linkages (degree of branching)
constitutes the basis for the characterization and definition of
the glucan components. Amylose is a relatively long, linear
polymer with few branches (1% a-(1,6)-linkages), whereas
amylopectin is a large, highly branched molecule, composed
of about 5% a-(1,6)-linkages. The structure of the intermediate
material falls in between those of amylose and amylopectin.
The molecular weight ranges of the amylose and amylopectin
are in the order of 105106 g mol 1 and 107109 g mol 1,
respectively. While the location of the amylose and its contri-
bution to the granular architecture are still largely unknown, it
is thought to be present primarily in the amorphous region of
the granules. Amylose chains that pass through both the crys-
talline and amorphous layers are referred to as tie chains.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00095-4
 CARBOHYDRATES | Starch: Starch Architecture and Structure
(a)
(c)
Figure 1 Light micrographs (1000 ) of maize (a), canna (b), mung bean (c), and Hylon VII maize starches (d).
Amylopectin Structure
Over the years, a number of models have been employed as a
means of visualizing the organization of amylopectin glucan
chains. Currently debated structural models include the tradi-
tional cluster model and the two-directional backbone model.
The traditional cluster model was originally proposed indepen-
dently by both French and Nikuni and was subsequently mod-
ified by Hizukuri. In this model, short adjacent linear
amylopectin chains are paired as double helices tightly regis-
tered in parallel alignment to form clusters, which are intercon-
nected through long chains. According to this model, long
amylopectin chains are part of the crystalline and amorphous
lamellae (Figure 2(a)). The model is based primarily on indirect
evidence drawn from the amylopectin unit chain profile.
Another argument that supports this model stems from the
molecular structure of lintners (acid-treated starch). This indi-
rect evidence suggests that the group of short chains builds up
the crystalline region, whereas long chains interconnect the
clusters. In the two-directional backbone model, the direction
of the short chains in the clusters is perpendicular to the direc-
tion of the long chains, which are present mainly in the amor-
phous lamellae. According to this model, long chains do not
pass through the crystalline region (Figure 2(b)). Interestingly,
the two-directional backbone model can also be defended based
on the indirect evidence and arguments put forward for the
traditional model. Unfortunately, no direct experimental verifi-
cation yet exits to enable the validity of either model to be tested,
and none of the amylopectin models developed are based on
the function of isozymes during starch biosynthesis.
191
(b)
(d)
The molecular structure of amylopectin from individual
starches can be differentiated according to both the unit
chain profile and the organization of the internal chains. The
unit chain profile describes the chain length (CL) distribution
pattern, or average composition, of the chains in the whole
amylopectin molecule, whereas the organization of the inter-
nal chains primarily identifies the nature of the branching
pattern.
Amylopectin Unit Chain Profile
After hydrolyzation of the a-(1,6)-linkages using debranching
enzymes, isoamylase and pullulanase, the unit chain profile of
amylopectin can be obtained through high-performance size-
exclusion chromatography (HPSEC) or high-performance
anion-exchange chromatography (HPAEC). Unit chains are
broadly categorized into three major groups: A, B, and C.
A-chains carry no other chains and are linked with the rest of
the molecule by a-(1,6)-bonds, whereas B-chains carry one or
several other chains (A- and/or B-chains). A C-chain carries the
sole reducing end of each amylopectin molecule.
Hizukuri described the polymodal distribution pattern of
amylopectin based on the observed periodicity of the CL
and subdivided the B-chains accordingly into B1, B2, B3, and
B4 categories. These chains differ with respect to their length
and the number of clusters they span. B1-chains (with a degree
of polymerization (dp) 20 to dp 24) are short chains, which
build up the clusters in partnership with A-chains (dp 11 to dp
14), whereas B2- (dp 42 to dp 48), B3- (dp 69 to dp 75), and
B4- (dp > 75) chains are long B-chains (dp > 36). The numerals
192 CARBOHYDRATES | Starch: Starch Architecture and Structure
f
A
(a)
C Short
A f
Amorphous
backbone
(b) (long chains)
Figure 2 (a) Schematic representations of the cluster model (adapted from Robin JP, Mercier C, Charbonniere R, and Guilbot A (1974)
Lintnerized starches. Gel-filtration and enzymatic studies of insoluble residues from prolonged acid treatment of potato starch. Cereal Chemistry 51:
389406) and (b) the backbone model of amylopectin (reproduced from Chauhan F and Seetharaman K (2013) On the organization of chains in
amylopectin. Starch/Starke 65: 191199.).
associated with B-chains indicate the number of clusters
they span (Figure 3). According to the divisions obtained by
Hizukuri using HPSEC, the periodicity of the CL is approxi-
mately dp 27 to dp 28, and the average CL is dp 17 to dp 26.
Hanashiro et al. observed a shorter periodicity (dp 12) with
HPAEC and named the chain fractions fa- (dp 6 to dp 12), fb1-
(dp 13 to dp 24), fb2- (dp 25 to dp 36), and fb3- (dp > 36)
chains, which presumably represent A-, B1-, B2-, and B3-
chains, respectively. Using this division, B2-chains are consid-
ered a subdivision of the B1-chains categorized by Hizukuri.
The very shortest amylopectin chains consist of six glucose
residues. The limit dextrin profile of amylopectin reveals that
A-chains (fa-chains) overlap with B1-chains (fb1-chains), so
the fa fraction therefore represents only part of the A-chains.
A-chains consist of a subgroup of chains called fingerprint
A-chains (dp 6 to dp 8), which are too short to form double
helices and can cause structural defects within the crystalline
lamellae. On a molar basis, the ratio of short to long unit
chains varies from 4 to 19, depending on the starch source.
Although the unit chain profile is useful for describing many of
the structural features of the amylopectin molecule, it fails to
explain how glucan chains are organized within the amylopec-
tin molecule.
Organization of Amylopectin Internal Chains
The organization of the unit chains in amylopectin determines
the unique architecture of the starch granule. Unit chains can
be categorized as either external or internal chains. External
chains are composed of A-chains and a portion of the B-chains.
The AB-chain segment that has branching points is designated
an internal chain. External chains participate in the formation
Reducing
end
Cluster
(short
chains)
Long chain
chains
Reducing
end
of the crystalline lamellae, whereas internal chains are present
primarily in the amorphous lamellae. The internal unit chain
profile can be obtained from f,b-limit dextrins by removing
the external amylopectin chains using the exo-acting enzymes
phosphorylase and b-amylase. Since exo-acting enzymes can-
not bypass branching points, the resulting limit dextrin repre-
sents the internal amylopectin structure. In f,b-limit dextrins,
all A-chains and the external segments of B-chains appear as
maltosyl and glucosyl stubs, respectively. The characterization
of A- and B-chains from f,b-limit dextrin provides a relatively
accurate estimate of the ratio of A-chains to B-chains, which,
on a molar basis, is typically in the range of 0.81.5.
The internal amylopectin structure consists of two major
groups of B-chains: short B-chains (dp < 24) and long B-chains
(dp > 24). Short internal B-chains (BS chains) are subdivided
into fingerprint B-chains (Bfp) with dp values 7 and a major
group of short B-chains (BSmajor) with dp values from 8 to 25.
Long internal B-chains are classified as B2-chains (dp 26 to dp
50) or B3-chains (dp > 50). Short B-chains are part of clusters,
and long internal B-chains function as cluster-connecting
chains. Based on the quantity and distribution pattern of the
short and long internal B-chains, amylopectin from different
botanical sources is one of four different types: type 1, type 2,
type 3, or type 4 (Figure 4). Type 1 amylopectin is character-
ized by large numbers of short internal chains, mainly BSmajor-
chains, with a broad distribution pattern, whereas type 4
amylopectin has a large number of long internal chains.
Types 2 and 3 represent intermediate distributions. Type 2
amylopectin exhibits a significant number of Bfp chains in
clusters. Based on this grouping, cereal starches are considered
to be types 1 and 2, and root and tuber starches to be primarily
types 3 and 4.
 CARBOHYDRATES | Starch: Starch Architecture and Structure 193

Reducing
A-chain B2-chain B3-chain B1-chain C-chain
end

Dp 1216

Figure 3 Chain categories of amylopectin. Adapted from Hizukuri S (1986) Polymodal distribution of the chain lengths of amylopectins, and its
significance. Carbohydrate Research 147: 342347.

Bfp BSmajor B2 B3

4
Carbohydrates (wt %)

0
0 20 40 60 80 100 120
Degree of polymerization (DP)

Type 1 (A-and C-crystalline): e.g., barley, rye, oat, Andean yam

Type 2 (A-and C-crystalline): e.g., maize, rice, sago, kudzu
Type 3 (A-and C-crystalline): e.g., cassava, arrowroot, mung bean
Type 4 (B-crystalline): e.g., potato, canna, lesser yam

Figure 4 Internal unit chain profile of debranched f,b-limit dextrins from four different types of amylopectin, as obtained by high-performance
anion-exchange chromatography (HPAEC). Adapted from Bertoft E, Piyachomkwan K, Chatakanonda P, and Sriroth K (2008) Internal unit chain
composition in amylopectins. Carbohydrate Polymers 74: 527543, with permission.

The cluster structure of amylopectin is examined primarily
with the use of enzymatic methods. Clusters, or macro a-limit
dextrins, are isolated from amylopectin using limited hydroly-
sis with a-amylase. These clusters are defined as groups of short
chains separated by internal chain segments with dp values
< 9. This definition is somewhat arbitrary because the hydro-
lysis pattern is dependent on the source of the a-amylase and
its kinetics as well as on experimental conditions. Clusters
from different types of amylopectin differ with respect to size
and CL and are made up of tightly branched units, known as
building blocks. Building blocks are micro a-limit dextrins,
which are released from clusters through extensive hydrolysis
with a-amylase. The size of the building block is determined
based on the number of chains (2, 3, 4, or more) in the unit.
Adjacent building blocks within the cluster are connected by
interblock chain segments, which are in the range of 58
glucose residues. Computer simulation and crystalline
stability studies have revealed that the CL between adjacent
building blocks is one of the most critical parameters govern-
ing the alignment of the external chains and the crystalline
packing. The number of building blocks within the cluster,
the interblock chain length (IB-CL), and the cluster size and
the number of chains in the cluster and the length of the
external chains are all used to characterize different types of
amylopectin. A schematic model illustrating the internal struc-
ture of amylopectin is provided in Figure 5. Cereal starches
194 CARBOHYDRATES | Starch: Starch Architecture and Structure
External chains
Building blocks Inter-cluster
chains (DP > 9)
Figure 5 Schematic illustration of the building block structure of amylopectin.
generally have a higher number of building blocks, a shorter
IB-CL, and larger cluster sizes than tuber and root starches. The
two-directional backbone model of amylopectin is based pri-
marily on the structure of experimentally isolated clusters and
building blocks.
Structural Polymorphism of Amylopectin
The double helices of amylopectin in the crystalline lamellae
are organized into different types of polymorphs. Starch gran-
ules exhibit two main types of polymorphic patterns: A-type
and B-type. They differ primarily with respect to packing
arrangement and degree of hydration. The A-type polymorph
is packed with monoclinic unit cells and consists of four H2O
molecules/unit cell, whereas the B-type has a hexagonal unit
with 36 H2O molecules/unit cell (Figure 6).
Polymorphic patterns can be characterized with the use of
wide-angle x-ray scattering (WAXS). Most starches derived
from cereals exhibit an A-type polymorphic pattern with char-
acteristic main peaks at 15 2y and 23 2y and an unresolved
doublet at around 17 2y and 18 2y. Characteristic of tuber, ae
maize, and retrograded starches, the B-type polymorphic pat-
tern shows primary reflection intensities at 5.5 2y, 1415 2y,
and 17 2y, as well as a well-resolved doublet centered at 22
2y24 2y. Root, pulse, and some fruit and stem starches
exhibit a C-type polymorphic pattern, which is a mixture of
A- and B-crystalline types (Figure 7). The type of polymorphic
pattern is thought to be related to the amylopectin CL (A-type
CL < 20 and B-type CL  22). Interestingly, a B-type polymorph
has also been observed in some starches (e.g., Arabidopsis leaf
and immature wheat grain) that have shorter amylopectin CLs
than B-type endosperm starches. This finding suggests that the
polymorphic patterns of starches cannot be explained based
solely on the average CL of the amylopectin.
Amylose Structure
Starches generally contain both linear and branched fractions of
amylose. Incomplete b-amylolysis indicates the presence of
branched fractions, which consist of 517 chains linked by
a-(1,6)-linkages. The conformation of amylose in an aqueous
solution is highly speculative. Since its random coil conforma-
tion is unstable, amylose tends to form single helices with guest
molecules (lipid, iodine, alcohols, and dimethyl sulfoxide
Interblock
chains length
(IB-CL)
B-type A-type
Figure 6 Unit cell structure of B- and A-type starches. Each green circle
represents the upper view of the double helix; the red dots indicate
water molecules. Adapted from Imberty A, Chanzy H, Perez S, Buleon A,
and Tran V (1988) The double-helical nature of the crystalline part of
A-starch. Journal of Molecular Biology 201: 365378.
(DMSO)) or double helices with adjacent chains in the absence
of complexing agents. Single-helix amylose is referred to as
V-amylose and is characterized by six glucose units per turn.
However, the helical structure and number of glucose units per
turn are controlled by the size of the inclusion complex. In
WAXS pattern, a peak centered at 20 2y generally reflects
V-type crystallinity. Reflection intensities of 7.5 2y and 13 2y
can also be considered indicative of a single-helix structure.
Double helices of amylose are found in A- or B-polymorphic
form and are similar to the crystallites formed by amylopectin.
Semicrystalline Structure of Starch Granules
Starch is known to have a hierarchically ordered structure with
characteristic periodicity. Its levels of organization are granule,
growth ring, blocklet, lamella, and glucan chains (Figure 8).
Each level has been characterized and defined with the use of a
variety of imaging and analytic techniques. However, the struc-
tural connections between these levels of order are still far from
completely understood.
The organization of glucan chains at different levels of order
determines the architecture of starch granules, which consists
of alternating amorphous and semicrystalline growth rings that
develop outward from the hilum by apposition. The thickness
of the growth rings generally varies from 120 to 400 nm. The
number and thickness of growth rings formed during granule
development are dependent on the botanical source of the
starch. Growth ring formation is thought to be subject to the
diurnal cycle, with semicrystalline rings being deposited during
 CARBOHYDRATES | Starch: Starch Architecture and Structure 195

A-type
B-type
C-type

0 5 10 15 20 25 30 35 40
2q ()
Figure 7 Wide-angle x-ray scattering patterns of A-, B-, and C-type starches.

Amorphous
background

Crystalline
lamella (C)
Amorphous
lamella (A)
9 nm

Maize/HCl/2.2M/ 80% hydrolysis

Cluster
Starch granule Growth rings Blocklets
(1 100 mm) (120 400 nm) (20 100 nm)

Double helix

C 2.1 nm
5 6 nm
9 nm

A
3 4 nm

Figure 8 Schematic representation of the structural levels of starch.

the daytime and amorphous rings being formed during the
night. However, whereas constant light treatment has been
shown to affect ring formation in barley and wheat, it has no
effect on ring formation in potatoes. While this finding sug-
gests that a circadian rhythm related to biological mechanisms
influences ring formation, no definite explanation has yet been
established for the occurrence of alternating zones of growth
rings in starch granules.
While the detailed structure of amorphous growth rings is
still to be determined, such rings are thought to consist of
amylose and disordered amylopectin. A semicrystalline growth
ring consists of a number of alternating crystalline and amor-
phous lamellae with a repeat distance of 8.511 nm. Amor-
phous lamellae are thought to contain amylose plus long
chains and branching points of amylopectin. Crystalline lamel-
lae are formed primarily from the double helices of short
amylopectin chains. Each double helix consists of six anhydro-
glucopyranose units per turn with a pitch height of 2.1 nm2.
The birefringence pattern of starch granules revealed under
polarized light microscopy is attributable to their
196 CARBOHYDRATES | Starch: Starch Architecture and Structure
(a)
Figure 9 Bright-field microscopy (a) and polarized microscopy (b) of iodine-treated waxy potato starch granules.
semicrystalline nature, and the distinctive Maltese cross indi-
cates that the ordered amylopectin chains in the crystalline
lamellae are radially oriented (Figure 9).
Images obtained using high-resolution scanning electron
and atomic force microscopy indicate another level of organi-
zation called a blocklet, which appears to be present in both
semicrystalline and amorphous growth rings. Blocklets are
more or less spherical, with sizes varying from 20 to 100 nm,
depending on the location of the granule and the botanical
source. Potato starch blocklets are generally larger than those
in cereal starches. The presence of the larger blocklets, coupled
with its semicrystalline nature, is commonly linked to the
relative resistance to enzymatic hydrolysis of the potato starch
granule. According to the blocklet concept, growth rings are
organized in the form of blocklets that occur in a semi-
crystalline growth ring consisting of a stack of lamellae. How-
ever, the exact nature of both the blocklet and its relation with
the rest of the structural levels of a starch granule remains open
to speculation.
Conclusion
In starch granules, amylopectin chains are organized into a
hierarchical structure. Although existing sophisticated imaging
and analytic techniques have enabled the observation of
different levels of order in starch granules, a substantial
knowledge gap remains with respect to understanding how
glucan chains are arranged in amylopectin and how the
structural levels are interconnected. The crystalline lamellae
and unit CL distribution of amylopectin have been more
extensively analyzed than other structural features. However,
the organization of glucan chains cannot be explained based
on the average composition of the unit chains. Detailed anal-
ysis of the internal amylopectin structure is therefore of
paramount importance for providing insight into the crystal-
line stability, the polymorphic pattern, and the architecture
of the building blocks, which are considered to be the funda-
mental structural unit in amylopectin. Enhanced knowledge
of the granular architecture will provide additional opportuni-
ties for uncovering the processes that govern the initiation
of starch synthesis, the control mechanism involved in
(b)
biosynthetic enzymes, and the structurefunction relationship
of starch in a variety of applications.
Exercises for Revision
How does amylopectin build up the semicrystalline
lamellae?
Define the structural parameters of amylopectin.
Explain how a,b-limit dextrin has improved the unit chain
analysis of amylopectin.
How does the internal structure of amylopectin determine
the glucan chain alignment in crystalline lamellae?
Exercises for Readers to Explore the Topic Further
How can the validity of the proposed amylopectin models
be tested?
Does the nature of the branching pattern influence the
polymorphic pattern?
Does amylose influence the assembly of crystalline
lamellae?
How does the molecular structure of amylopectin in the
semicrystalline region of starch granules differ from the
amylopectin in the amorphous region?
Do blocklets build up the super helical structure of starch
granules?
See also: Carbohydrates: Starch: Analysis of Quality; Starch:
Chemistry; Starch: Synthesis; Genetics of Grains: Maize: Other
Maize Mutants; Processing of Grains: Starch: Modification; Starch:
Uses of Native Starch.
Acknowledgment
The authors thank Dr Eric Bertoft for his critical review of this
article.
 CARBOHYDRATES | Starch: Starch Architecture and Structure
Further Reading
Bertoft E (2013) On the building block and backbone concepts of amylopectin structure.
Cereal Chemistry 90: 294311.
Bertoft E, Piyachomkwan K, Chatakanonda P, and Sriroth K (2008) Internal unit chain
composition in amylopectins. Carbohydrate Polymers 74: 527543.
Buleon A, Colonna P, Planchot V, and Ball S (1998) Starch granules: Structure and
biosynthesis. International Journal of Biological Macromolecules 23: 85112.
Chauhan F and Seetharaman K (2013) On the organization of chains in amylopectin.
Starch/Starke 65: 191199.
French D (1984) Organization of starch granules. In: Whistler RL, BeMiller JN, and
Paschall EF (eds.) Starch: Chemistry and Technology, 2nd edn., pp. 183207.
San Diego, CA: Academic Press.
Hanashiro I, Abe J, and Hizukuri S (1996) A periodic distribution of the chain length
of amylopectin as revealed by high-performance anion-exchange chromatography.
Carbohydrate Research 283: 151159.
Hizukuri S (1986) Polymodal distribution of the chain lengths of amylopectins, and its
significance. Carbohydrate Research 147: 342347.
Imberty A, Chanzy H, Perez S, Buleon A, and Tran V (1988) The double-helical
nature of the crystalline part of A-starch. Journal of Molecular Biology
201: 365378.
197
Nikuni Z (1978) Studies on starch granules. Starke 30: 105111.
OSullivan AC and Perez S (1999) The relationship between internal chain length
of amylopectin and crystallinity in starch. Biopolymers 50: 381390.
Perez S and Bertoft E (2010) The molecular structures of starch components and
their contribution to the architecture of starch granules: A comprehensive review.
Starch 62: 389420.
Robin JP, Mercier C, Charbonniere R, and Guilbot A (1974) Lintnerized starches.
Gel-filtration and enzymatic studies of insoluble residues from prolonged acid
treatment of potato starch. Cereal Chemistry 51: 389406.
Shannon JC and Garwood DI (2009) Genetics and physiology of starch development.
In: Whistler RL and BeMiller JN (eds.) Starch: Chemistry and Technology, 3rd edn.,
pp. 2586. San Diego, CA: Academic Press.
Thompson DB (2000) On the non-random nature of amylopectin branching.
Carbohydrate Polymers 43: 223239.
Vamadevan V, Bertoft E, and Seetharaman K (2013) On the importance of organization
of glucan chains on thermal properties of starch. Carbohydrate Polymers
92: 16531659.
Zobel HF (1988) Starch crystal transformations and their industrial importance. Starch
40: 17.
 Starch: Analysis of Quality
A Gunaratne and H Corke, The University of Hong Kong, Hong Kong, Peoples Republic of China
2016 Elsevier Ltd. All rights reserved.
This article is reproduced from the previous edition, volume 3, pp. 202212, 2004, Elsevier Ltd with revisions made by the editor.
Topic Highlights
Starch as the main constituent of food grains and the reason
for many of the functional properties of grains.
Starch occurs as granules in the endosperm of cereal grains;
granule shape is distinctive for the species.
The functional properties of starch are affected by botanical
source, structure and physicochemical properties, and pre-
treatments during processing.
The two major compounds of starch are amylose (mainly
linear polymers of a(14)-linked glucose) and amylopectin
(usually the major component, being highly branched
polymers).
Amylose forms a distinctive blue color complex with
iodine. Better analysis is provided by high-performance
size-exclusion chromatography (HPSEC) and various
other methods.
Near-infrared (IR)-reflectance analysis can be used to deter-
mine amylose content rapidly for mutant lines that differ
significantly in amylose:amylopectin ratio.
Direct testing of starch function involves the pasting test
whereby viscosity changes are monitored for a starch sus-
pension heated in excess water, using equipment such as
the Viscoamylogram or the Rapid ViscoAnalyzer.
Starch retrogradation is the key physical change associated
with bread staling.
Learning Objectives
Understand the chemical properties of starch that affect its
functional utilization.
Understand how and when to apply the range of testing
procedures for starch chemistry and function.
Introduction
Starch is the main polysaccharide produced in photosynthetic
tissues. Recently, there has been a growing demand for starch
in both food and nonfood industries because of its wide array
of applicability in diverse systems. Functionality is the key for
the industrial utilization of starch. Functional properties can be
defined as the characteristics that govern the behavior of a food
component during processing, storage, and preparation. Fac-
tors such as botanical source, structure and physicochemical
properties, and pretreatments of starch directly affect func-
tional properties of starch. Analysis of starch-related function-
ality should provide a clear picture of the relevant property that
will allow better use of starch in food and nonfood systems, for
example, to maintain consistent quality and functionality of
starch-based products in every batch, otherwise the product
198 Encyclopedia of Food Grains, Second Edition
will not survive in the competitive market. Methods that are
implemented for the analysis of starch functional properties
should be simple and fast, technically feasible, nondestructive,
and able to provide reproducible results with high accuracy.
Although several methods have been developed recently to
monitor some of the functional properties, individual methods
are sometimes not capable of providing adequate data on all
the dimensions of a functional property, especially in the
determination of retrogradation and gelatinization. Therefore,
it is often worth applying at least two methods to get comple-
mentary data. This article aims to discuss some functional
properties of starch from the perspective of the applicability
of methods that are currently employed to determine those
properties.
Properties, Functions, and Analysis of Starch
Macromolecules
Amylose
The two major compounds of starch are amylose and amylo-
pectin. The two polymers are structurally different, the minor
component, which is linear, consisting mainly of a(14)-
linked glucose. However, in some starches, amylose chains
have shown a slight degree of branching. Incomplete conver-
sion to maltose by b-amylase suggests the presence of branched
points in the amylose polymer. The branched chains can also
act similar to unbranched chains. Amylose isolated from tuber
and root starches such as potato and cassava has a larger
molecular weight than that isolated from cereal starches such
as maize and wheat. Although amylose is a linear molecule, its
conformation tends to change due to the large number of
hydroxyl groups that can produce higher hydrogen bonding
capability with strong internal forces. The conformation of
amylose has been the subject of controversy and has been
shown to vary from helical to an interrupted helix, to a random
coil. In alkaline solutions (KOH) and dimethyl sulfoxide
(DMSO), amylose probably has an expanded coil conforma-
tion, while in water and neutral aqueous potassium chloride
solutions, it is a random coil with short, loose helical segments.
Although helical conformation is common in amylose, double
helices form when different helices are packed together. The
helical arrangement of amylose forms a hydrophobic core
inside the amylose molecule, which permits complexing with
guest molecules such as lipids and iodine. The amount and
molecular characteristics of amylose have a significant influ-
ence on starch functionality. Higher amylose content decreases
the granular swelling power, whereas increasing amylose con-
centration decreases the gel stickiness but increases the gel
firmness. Reassociation of amylose chains on cooling of a
starch paste shrink the starch gel resulting in water
http://dx.doi.org/10.1016/B978-0-12-394437-5.00092-9
 CARBOHYDRATES | Starch: Analysis of Quality 199

Table 1 Some physical and chemical properties of common starches

Starch Granular size (mm) Granular shape Amylose (%) Swelling power at 95  C Solubility at 95  C Taste

Barley 235 Round, elliptical, lenticular 22 Low

Maize
Regular 525 Round, polygonal 26 24 25 Low
Waxy 525 Round, oval 1 64 23 Low
High amylose Round Up to 80 6 12 Low
Potato 15100 Egg-like, oyster indentations 22 1000 82 Slight
Rice 38 Polygonal clusters 17 19 Low
Rye 235 Elliptical, lenticular 23 Low
Sago 2060 Egg-like, some truncate forms 27 97 Low
Sorghum 525 Round, polygonal 26 22 22 Low
Tapioca 535 Round, oval, truncated on side 17 71 48 Fruity
Wheat 235 Round, elliptical, lenticular 25 21 41 Low
Oats 210 Polygonal, compound 27 Low

Adapted from Collado LS and Corke H (2003) Starch properties and functionalities. In: Kaletunc G and Breslauer KJ (eds.) Characterization of Cereals and Flours, pp. 473506.
New York: Marcel Dekker.

accumulation on an aging gel, which decreases the storage Table 2 Amylose content in maize mutant starches as estimated with
stability and usually the quality of an affected starch-based size exclusion chromatography (SEC), Concanavalin (Con. A), differential
food product. scanning calorimetry (DSC), and iodine binding capacity (IBC)

Amylose (%)
Methods used in amylose estimation
Starches SEC Con. A DSC IBC
Conventional quantitative determination of amylose is based
on its ability to form a deep blue color complex with iodine. ae wx nd 7 nd 14
Amylose content of starch therefore can be quantified either by Du 27 45 55 45
spectrophotometric means or by potentiometric titration. su2 24 50 58 45
Table 1 shows amylose content of some common starches. du su2 34 58 66 60
However, interference from amylopectin and other intermedi- ae du 30 56 64 56
ate materials in the blue color formation reaction biases the Ae 54 63 60 63
estimation of actual content of amylose. Amylopectiniodine nd: not detected.
complex is not stable due to shorter unit chain length of Source: Planchot V, Gerard C, Bertoft E, and Colonna P (2001) An approach to
amylopectin, but long chains of amylopectin, e.g., long B structural analysis of granules using genetically modified starches. In: Barsby TL,
chain of waxy maize amylopectin can bind with iodine in a Donald AM, and Frazier PJ (eds.) Advances in Structure and Function, pp. 104128.
similar way to amylose. This would lead to an overestimation. UK: The Royal Society of Chemistry.
Despite shortcomings such as this, and the time-consuming
nature of the assay, the amyloseiodine reaction is still the
basis of the most widely used method to determine amylose component and the resulting supernatant after centrifugation
content of starch because of its accuracy, reproducibility, and contains only the amylose component. However, a compara-
easy operation. However, the excessive time needed limits its tive study of size exclusion chromatography (SEC), differential-
use in quality control applications. Instead of conventional scanning calorimetry, iodine-binding capacity (IB), and
thermal dissociation of amylose in DMSO, low-temperature concanavalin-A, where SEC was used as the reference method,
gelatinization in CaClB2B could be employed to reduce the has revealed that IB, differential-scanning calorimetry (DSC),
time requirement. High performance size exclusion chroma- and concanavalin-A resulted in an overestimation in the
tography (HPSEC), that has shown a good correlation with determination of amylose content of mutant maize starches
blue value measurements of amylose content, can be employed (Table 2). As other alternative methods to the conventional
to separate starch and de-branched starch. Ability to directly iodineamylose complex formation, spectroscopic methods
monitor the effect of de-branching on the molecular size have been developed recently. These techniques provide advan-
distribution of starch and high-molecular weight linear amy- tages such as ease of sample preparation, speed, and applica-
lose content from this technique provides the estimation of bility to online monitoring in the process control situation
long-chain amylopectin chains that contribute to the apparent compared with other techniques. Near-infrared (IR)-reflectance
amylose content. Recently another technique that is able to analysis has been successfully introduced in determining the
eliminate the effect of long-chain amylopectin in the estimation amylose content of waxy, normal, and high amylose maize,
of amylose content has been introduced. In this method, starch unground brown or milled rice, and whole grain maize sam-
completely dissolved in DMSO is subjected to de-fatting with ples. Raman spectroscopy on the other hand is becoming more
ethanol. The precipitate after de-fatting is dissolved in acetate/ popular as a quantitative analytical technique in the food
salt solution. A special reaction mixture (concanavalin-A) is industry and has shown potential applicability in the estima-
then added to the solution to precipitate the amylopectin tion of amylose content in maize starches.
200 CARBOHYDRATES | Starch: Analysis of Quality
Amylopectin
Amylopectin, the highly branched molecule, is usually the
major component in the starch granule with a(14)-linked
glucose linear chains and a(16)-linked branch points. Crys-
talline domains of the starch granules are due to the clustered
branches of amylopectin chains that are packed together,
whereas the free amylose, amylose complexed with lipids,
and branch points of the amylopectin are found in the amor-
phous region. Alternative arrangement of crystalline and amor-
phous region was proposed for the semicrystalline starch
granule. However, there is no clear demarcation between
amorphous and crystalline regions. Crystalline region is less
susceptible to enzymatic hydrolysis, water penetration, and
other chemical reactions than amorphous region. Amylopectin
has a lesser tendency to gelation, retrogradation, and syneresis
because of the branched structure. The amount of amylopectin
varies among different starches. Waxy varieties contain almost
100% amylopectin. The extent of functional characteristics of
starch (viscosity, gelatinization, solubility, texture, gel stability,
retrogradation, shear resistance) are directly affected by the
amylose/amylopectin ratio. Although amylopectin is the
major component of the starch granule, there is no convenient
method developed for direct estimation of amylopectin and
studies on amylopectin are dependent on development of
enzymatic and instrumental methods. Average structural prop-
erties of whole molecule and impact of internal structure
of clusters on crystallization have been studied by means of
enzymatic and SEC. It has been shown that the combined
application of preparative and analytical size exclusion chro-
matography with multiple detection, precipitation techniques,
and enzymatic de-branching provides the most detailed
analytical insight on the microstructural properties of
amylopectin.
Function Related to Morphological Characteristics
of Starch Granules
Granules are the basic physical structural unit of starch. The
size, shape, and other morphological characters of starch gran-
ules are extremely diverse (Table 1). The shape of the starch
granule can be spherical, disk, polygonal, or elongated. In
general, cereal starches have smaller granules compared to
tuber and root starches. Most tuber and root starch granules
have a simple size distribution. In some starches, such as barley
and wheat, two different granular size populations exist, i.e., a
bimodal distribution. Granular size and morphology have
recently received more attention, e.g., the size of granules is
important in determining taste and mouth feel of some starch-
based fat mimetics. Some specific industrial applications of
starch are related to size and the size distribution of starch
granules. For example, the small size of rice starch granules
makes it highly suitable for laundry use. Most of the physico-
chemical properties of native starch are highly correlated with
the size of the starch granules. Larger size granules tend to swell
more than smaller granules, whereas smaller size granules are
more susceptible to enzymatic hydrolysis. The outer surface of
starch granules also plays a key role in many applications of
starch. Electron microscopic images have revealed that tuber
and root starches show no sign of any pores on the granular
surface. However, pores are present on the surface of maize,
sorghum, and millet starch granules. Pores on granule surfaces
increase accessibility of a-amylase to the granule interior result-
ing in increased enzymatic hydrolysis. Microscopic analysis
provides information on surface features and size of starch
granules.
Characterization and Analysis of Starch Thermal
Properties
Gelatinization
Gelatinization is one of the most important processes affecting
starch. It occurs when starch is heated in excess water, when
irreversible granular swelling, native crystallites melting, loss of
granular order, loss of birefringence, and starch solubilization
take place. The extent of these changes depends on type of
starch, starch concentration, temperature, presence of other
solutes, and shear applied during the gelatinization process.
As a result of the above changes during gelatinization, the
starch paste develops viscosity, the basis of most technological
usefulness of starch as an ingredient in food and nonfood
applications. In the majority of food applications, starch func-
tions (e.g., imparts texture on consumption) in a gelatinized
form. Several concepts have been proposed to explain the
gelatinization process. Granular swelling acts primarily as a
driver to destabilize the starch crystallites. Recently, a model
was proposed to explain the phenomena involved in gelatini-
zation and hydration of starch based on the side-chain liquid
crystalline. In this model, lamellae in starch are considered in
terms of three components (1) backbone, (2) side-chain, and
(3) double helices. The degree of mobility of these three com-
ponents, coupled with the helixcoil transition, give starch its
distinctive properties in gelatinization. Gelatinization is influ-
enced by many factors such as, botanical source of starch, water
content, and added solutes. Several methods have been pro-
posed to study the effect of water content on thermal stability.
Depending on the water content available during gelatiniza-
tion, changes in endothermic transition have been detected in
terms of number and the position of the differential scanning
calorimetric endotherms. Only one endotherm is detectable at
higher levels of moisture content while two endothermic tran-
sitions are reported at the low-moisture-content level for some
starches where low temperature and higher temperature endo-
therms are designated as G1 and M1 respectively (Figure 1). In
addition to two endothermic transactions at lower moisture
content, peaks are shifted to higher temperature, indicating
need for higher thermal energy to melt the starch crystallites
at low moisture content. If there is sufficient water available in
gelatinization, then crystallites melt cooperatively, resulting
in a single endotherm. Two endothermic transition peaks in
insufficient water could be attributed to the differences in the
stability of starch crystallites, which melt over different tem-
peratures, less stable crystallites melt first and the others are
next causing the second endotherm. Redistribution of water
between crystalline domain and gelatinized starch could also
result in two endothermic transitions, in which the first endo-
therm could be attributed to melting of starch crystals in excess
water. Much research has been conducted to investigate the
 CARBOHYDRATES | Starch: Analysis of Quality 201

Temperature (C)

To Tc
Hcx
M2
Hgel

Endothermic heat flow

Tcx

G M1

Tm

Figure 1 Schematic representation of DSC thermograms obtained for wheat starch heating. (Eliasson A-C (2003) Utilization of thermal properties for
understanding baking and salting processes. In: Kaletunc G and Breslauer KJ (eds.) Characterization of Cereals and Flours, pp. 65115. New York:
Marcel Dekker.)

Table 3 Gelatinization parameters of some starches measured by differential scanning calorimetry

Starch Starchwater ratio To Tp Tc To Tc DH

Wheat 1:3 57.0 62.0 67.0 10.0 9.7

Maize normal 1:3 65.3 71.3 80.9 15.6 11.0
Maize waxy 1:3 62.9 72.8 84.3 21.4 13.6
Potato 1:3 59.6 66.3 76.0 16.4 16.3
Cassava 1:3 63.0 71.5 81.5 18.7 12.3
Sweet potato 1:3 60.0 69.0 82.5 22.5 7.1
Taro 1:3 76.8 83.0 95.2 18.4 14.5
True yam (Dioscorea) 1:3 75.0 80.0 90.2 16.5 17.8

To onset temperature ( C); Tp peak temperature ( C); Tc conclusion temperature ( C). DH enthalpy (J g 1).
Sources: Gunaratne A and Hoover R (2002) Effect of heat-moisture treatment on the structure and physicochemical properties of tuber and root starches. Carbohydrate Polymers 49:
425437; and Hoover R, Vasanthan T, Senanayake NJ, and Martin AM (1994) The effects of defatting and heat-moisture treatment on the retrogradation of starch gels from wheat, oat,
potato, and lentil. Carbohydrate Research 261: 1324.

influence of added solutes such as sugars, emulsifiers, and
electrolytes (such as sodium chloride), which are widely
applied in food industry on starch behavior. Sugars have
shown to increase the gelatinization temperature but not to
affect endothermic heat absorption. Quality of various food
products, bread and cakes, thickening and gelling of sauce, pie
filling, and extrusion of cereals are highly dependent on
starchwater interaction. Increased gelatinization, when sugars
are added to starch paste could be due to decrease of water
availability for the gelatinization, binding water to sugar mol-
ecules. However, it was reported that sugar acts as an antiplas-
ticizer rather than gelatinizer through the effect on water-
binding capability. Increased sugar concentration decreases
the plasticization effect requiring more heat energy to achieve
the gelatinization. The extent of the influence of sugars on
gelatinization, swelling, and viscosity differs depending on
the type of sugars and salts used when compared on a molar
basis. Monosaccharides are less effective than disaccharides
except maltose; among disaccharides, sucrose is more effective.
Methods in the analysis of starch gelatinization
There are several methods to analyze the gelatinization process,
such as DSC, Kofler hot-stage microscopy, light microscopy,
electron microscopy, X-ray crystallography, enzymatic analysis,
nuclear magnetic resonance (NMR), pulsed nuclear magnetic
resonance, small angle X-ray scattering, and small angle neu-
tron scattering, Brabender viscoamylography (BV), and Rapid
Visco Analysis. DSC, the most widely used method, measures
the dissociation parameters, TBoB (onset), TBpB (peak), TBcB
(conclusion), and DH (endothermic heat absorption), of
starch crystallites in gelatinization (Figure 1). DH is the area
below the transition endotherm. Gelatinization parameters of
starches from different botanical sources measured by DSC are
presented in Table 3. In addition, DSC can be used to study the
glass transition temperatures of various starchy food products.
Gelatinization temperatures are influenced more by the gran-
ular architecture than the amyloseamylopectin ratio and
depend on the degree of starch crystallite perfection, whereas
endothermic heat absorption reflects the quality and amount
of starch crystallites. Analysis of gelatinization parameters of
starch using DSC was first reported in 1971. Gelatinization
parameters measured by DSC can be affected by sample prep-
aration, baseline application, pan selection, method of adding
water, sample mass, heating rate, and moisture equilibration
time. Thermal behavior of amyloselipid complex as well as
formation of amyloselipid inclusion with various molecules
such as fatty acids and emulsifiers can also be detected by DSC.
Characterization of amyloselipid complex in cereal starches
and influence of amyloselipid complex on starch properties
are well documented. Some of the internal lipids in cereal
202 CARBOHYDRATES | Starch: Analysis of Quality

starch is thought to be complexed with amylose to form the
amyloselipid complex, which is dissociated at higher temper-
ature (M2 endotherm in Figure 1) than that of normal starch
crystallites (Figure 1 and Table 4). Although DSC is widely
used to study gelatinization, the small sample size used can
limit its real applications. In a comparative study of maize
starch gelatinization with DSC and NMR, it was reported that
NMR could provide much better performance in the analysis of
gelatinization for larger samples than DSC. Gelatinization tem-
perature of Peruvian carrot, potato, and maize starch measured
by Rapid Visco Analyzer (RVA) and BV, and DSC have shown
different values for different methods. This indicates the need
for multiple analytical techniques for a meaningful under-
standing of gelatinization along with careful specification
(and preferably standardization) of experimental conditions
used. Table 5 shows the differences in gelatinization parame-
ters measured by different techniques for the same starch.
Pasting properties
Starch heated in excess water undergoes various changes as a
result of heat and moisture transfer. Gelatinization and pasting
occur in the same system and have often been used to describe
all the changes that occur. Gelatinization may be used to refer
to early changes whereas pasting includes later changes. Starch
gelatinization is defined as the collapse of the starch granule
manifested in irreversible melting, loss of birefringence, and
starch solubilization. The point of gelatinization, and the range
over which it occurs is governed by starch concentration,
method of observation, granular type, and heterogeneity. Past-
ing is defined as the phenomena following gelatinization in
the dissolution of starch, involving granular swelling, exuda-
tion of molecular components from the granule, and eventu-
ally total disruption of the granules. The schematic
representation of granular changes and viscosity development
that occurs during pasting is shown in Figure 2.
Measurement of pasting properties
The BV is the most widely established method for determining
pasting properties, although it has some methodological and
geometrical shortcomings. This apparatus measures the devel-
opment of viscosity when a starchwater suspension is sub-
jected to a programmed heating and cooling cycle under a
shear force. The temperature at which viscosity begins to
increase is termed pasting temperature. With further heating,
granular swelling increases the viscosity to reach a peak. Fur-
ther heating at elevated temperature under shear force tends to
disintegrate the swollen granules resulting in decrease of vis-
cosity. Upon cooling the starch paste in the next stage, there is a
tendency, mainly due to amylose chain reassociation, to
increase viscosity. Therefore, a typical pasting profile exhibits
Viscosity

Table 4 DSC parameters of amyloselipid complex of some cereal

starches

Starch Condition Tcx DHcx

Wheat 50% water 110.1 1.4

Rye 50% water 107.8 0.8
Barley 50% water 110.3 1.8 Time/temperature
High-amylose barley 50% water 110.8 2.8
Figure 2 Schematic representation of granular changes and viscosity
Tcx (peak temperature); DHcx (transition enthalpy). development during starch pasting (Daniel JR and Whistler RL (1985)
Source: Fredriksson H, Silverio J, Andersson R, Eliasson A-C, and Aman P (1998) The Principal changes in starch during food processing. In: Richardson
influence of amylose and amylopectin characteristics on gelatinization and TR and Finley JW (eds.) Chemical Changes in Food During Processing,
retrogradation properties of different starches. Carbohydrate Polymers 35: 119134. pp. 305326. Westport, CT: Avi Publishing Co.).

Table 5 Gelatinization temperature range of some starches as measured with different techniques: DSC, NMR, BV, and RVA

Gelatinization temperature range

Starch Starchwater ratio DSC NMR BVa RVAb

Maize 1:2 58.179.6 5967

Waxy maize 1:2 56.481.3 6067
Potato 1:2 49.273.9 5360 7686 7586
Cocoyam 1:2 7487 7895 8595
Peruvian carrot 1:2 5673 6295 6895
a
Starch concentration (4%) except Peruvian carrot (6%).
b
Starch concentration (8%).
Sources: Perez EE, Breene WM, and Bahnassey YA (1998) Gelatinization profiles of cassava, sagu, and arrowroot native starches as measured with different thermal and mechanical
methods. Starch/Starke 50: 1416 and Gonera A and Cornillon P (2002) Gelatinization of starch/gum/sugar systems studied by using DSC, NMR, and CSLM. Starch/Starke 54:
508516.
 CARBOHYDRATES | Starch: Analysis of Quality 203

three distinct viscosity developments, peak viscosity, hot paste
viscosity, and cold paste viscosity (Figure 3). Pasting curves
vary according to botanical source of the starch, starch concen-
tration, and the programmed heatingcooling cycle chosen.
Among the native starches, potato shows the highest peak
viscosity, and generally, low peak viscosity and higher setbacks
can be detected in normal cereal starches compared to tuber
and root starches. Waxy cereal starches, on the other hand,
behave more like the tuber and root starches (Figure 4).
According to the characteristics of pasting curves, starch has
been categorized into four groups as, high swelling starch
(potato, cassava, waxy cereals, ionic starch derivatives), mod-
erate swelling starch (normal cereal starch), restricted swelling
starch (cross-linked starch), and highly restricted starch
250
Temperature Cool paste viscosity
profile
200
Peak viscosity
Peak time
Setback
(starches >55% amylose). The above four different categories
of starch exhibit the following pasting properties respectively:
high peak followed by rapid and major thinning during cook-
ing; lower peak and much less thinning; no peak but maintain
high viscosity during cooking; and no swelling to gave a vis-
cous paste at normal concentration.
Because of some technical shortcomings with the BV, such
as large sample size requirement and inability to program the
temperature profiles, a more recent equipment, the RVA, has
become popular for analyzing pasting properties. The RVA
differs from the Brabender instrument due to rapid heating
rate and stronger mixing action. However, controlled heating
rate to 1.5  C min 1 in RVA provides similar results to those
observed in the BV. Comparative study of pasting properties of
cassava, sago (sagu), and arrowroot using Brabender and RVA
has shown that both techniques provide similar pasting pat-
terns (Table 6). Pasting curves of some of the starches deter-
100
mined by BV and RVA is presented in Figures 4 and 5.
80
Swelling and Solubility
Native starch granules are insoluble in water. Although small
amounts of water can be absorbed at room temperature, gran-
Temperature (C)

Breakdown
Viscosity (RVU)

150 60 ular swelling is limited in intact granules. During heating in

excess water, after the onset of gelatinization, granules begin to
swell rapidly, losing the polarization crosses. The extent of the
swelling power and solubility depends on the magnitude of the
100 Hot paste viscosity 40
starch chain interaction within the amorphous and crystalline
domain, size of the starch granules, amylose content, bound
lipids, starch damage, temperature, and molecular characteris-
50 20
tics and the amount of amyloseamylopectin ratio. Table 1
presents the swelling and solubility of some starches at 95  C.
High amylose content lowers the swelling whereas larger gran-
0 0 ules show greater swelling. The swelling power and solubility
0 3 6 9 12 15
provide information on the relative strength of bonding within
Time (min)
the granules.
Figure 3 Characteristics of a typical pasting curve.
Methods used to analyze starch swelling
Swelling volume, swelling power or swelling factor, all mea-
sure the extent of granular swelling at a given temperature,
Time when starch is heated in excess water. However, swelling factor
0 30 60 90 120 150 180 min
measures only the intergranular water content and hence
3000
reflects the true swelling of starch granules at a given temper-
ature. Swelling power measures the weight of swollen starch
2500
5% starch sediment relative to the initial starch dry weight, and is
Potato by weight expressed as the ratio of the wet sediment to the initial weight
2000
of the dry starch. Measuring the soluble carbohydrate that
dissolved in the supernatant either by oven drying or colori-
1500
metric method enables solubility, the ratio between the
Tapioca Maize amount of soluble carbohydrate to the initial weight of dry
Viscosity (BU)

1000
starch to be obtained. Swelling volume is a modified version in
Waxy maize
which programmed shaking of starch suspension is used
500
instead of stirring, and measurement is of the volume
Wheat expanded due to granular swelling. Swelling factor is the ratio
0
of volume of swollen starch granule to the volume of the dry
50 95 95 95 50 50 50 C
starch. Swelling factor measures only the intragranular water
Heat Hold Cool Hold
content. Calculation of the swelling factor is based on starch
Temperature weight adjusted to 10% moisture assuming the density of
Figure 4 Brabender Viscoamylogram of starches from different 1.4 mg ml 1. The basic principle of this method is the mea-
botanical sources. surement of the volume of water absorbed by starch granules
204 CARBOHYDRATES | Starch: Analysis of Quality

Table 6 Pasting properties of cassava, sago, and arrowroot as determined by BV and RVA

Starch Method GT PV FV at 95  C V at 50  C BD SB Consistency

a
Cassava BV 6890 50 40 40 10 10 0
RVAb 7390 23 9 17 14 6 8
Sago BV 6890 300 460 660 160 360 200
RVA 7395 60 22 37 38 23 15
Arrowroot BV 7590 150 150 170 0 20 20
RVA 7992 30 14 18 16 12 4

GT gelatinization temperature range; PV peak viscosity; FV final viscosity; V viscosity; BD breakdown; SB setback.
a
Starch concentration 4.3%.
b
Starch concentration 8%.
Source: Perez EE, Breene WM, and Bahnassey YA (1998) Gelatinization profiles of cassava, sagu, and arrowroot native starches as measured with different thermal and mechanical
methods. Starch/Starke 50: 1416.

Effect of retrogradation on quality of starch-based food

100
600 products
Retrogradation has desirable as well as undesirable effects on
quality of starch-based food products. Mostly it is undesirable.
Potato Bread staling is one of the main undesirable effects of retrogra-
450 80
dation. Although several factors contribute to bread staling, it
Temperature (C)
Viscosity (RVU)

Sweet potato has been found that retrogradation is the key physical change
associated with bread staling. Retrogradation sometimes aids
300 Maize 60 processing in some food products such as hardening of par-
boiled rice and to improve textural characteristics of certain
types of noodles.
150 40
Determination of starch retrogradation
Rice Several methods have been developed to determine starch
Wheat
0 retrogradation, because of its great influence on industry-
0 5 10 15 20 25 based food and nonfood products, such as DSC, X-ray analysis,
Time (min) rheological methods, and spectroscopic methods.

Figure 5 Pasting curves obtained for different starches from Rapid

Visco Analyzer.
heated in excess water, based on the observation that blue
dextran dye (molecular weight 2  10P6P) will dissolve in
supernatant and interstitial water. The advantages of this
method over the others are the ability to measure the true
swelling and the high accuracy.
Retrogradation
Reassociation of starch polymers via hydrogen bonding in
gelatinized starch on cooling is generally termed retrograda-
tion, and is time- and temperature-dependent. Starch gels tend
to undergo structural changes during storage as they are meta-
stable and nonequilibrium systems. Both amylose and amylo-
pectin are involved in retrogradation, where rapid amylose
aggregation causes a short-term development of starch gel
providing initial firmness. Branched amylopectin recrystalliza-
tion, particularly outer branches of amylopectin molecule, is
correlated with long-term development of starch gel. Several
factors influence starch retrogradation, such as, starch concen-
tration, storage temperature, initial heating temperature, chain
length distribution of amylopectin, molecular size of amylose,
lipids, and physical and chemical modification of starch.
Differential scanning calorimetry
Several studies on retrogradation behavior have been exam-
ined using DSC techniques. DSC is probably the best thermal
analysis method to study the starch-aging process for different
systems. However, dissociation parameters of stronger crystals
that formed due to the association of amylose are usually
difficult to detect using DSC, since those crystals dissociate at
higher temperature (>120  C). Therefore, DSC usually mea-
sures the melting of recrystallized amylopectin, that occurs at
the same temperature interval as gelatinization (Figure 6),
when reheating of retrograded starch gel, in which heat absorp-
tion, the area under the endotherm (DH), and transition tem-
peratures can be detected (Table 7). Crystals formed by the
association of amylose chains would be possible to detect
using DSC pans, that can withstand a higher temperature
range (Figure 7). Characteristic smaller endosperms were usu-
ally reported for the retrograded gel compared with the gelati-
nization endotherm of their native counterpart. However, it
was reported that retrograded starch crystals melt over a wide
range of temperature indicating the more heterogeneous crys-
tal perfection than those of native starch crystals. Melting tem-
perature indicates the perfection of recrystallized amylopectin,
the quality and heterogeneity.
Although DSC has some advantages especially that it is not
time consuming, has a wide range of applications, and mea-
sures directly, it has some weakness particularly its small
 CARBOHYDRATES | Starch: Analysis of Quality 205

Heating steps
Bread crumb
1

2
Wheat starch
Endothermic heat flow

Endothermic heat flow

Amylopectin 4

Amylose
0.05 mcal s1

40 60 80 100 60 100 140 180

Temperature (C)
Figure 6 Differential scanning calorimetric curves for different
retrograded food systems. (Eliasson A-C (2003) Utilization of thermal
properties for understanding baking and salting processes. In: Kaletunc
G and Breslauer KJ (eds.) Characterization of Cereals and Flours, pp.
65115. New York: Marcel Dekker.)
Temperature (C)
Figure 7 DSC curves obtained for amylose retrogradation. (Eliasson
A-C (2003) Utilization of thermal properties for understanding baking and
salting processes. In: Kaletunc G and Breslauer KJ (eds.) Characterization
of Cereals and Flours, pp. 65115. New York: Marcel Dekker.)

Table 8 X-ray pattern and crystallinity of different starches

Table 7 DSC characteristics of amylopectin retrogradation of wheat Starch X-ray pattern Crystallinity
and potato starch gel (1:1) under different storage conditions
Rice A 38
Starch Storage conditiona (To Tc) DH (J g 1
AMP) Oat A 33
Wheat A 36
Wheat 6/6 40.463.3 8.1
Rye A 34
6/30 48.564.7 7.7
Amylomaize B 1522
6/306/30 49.365.4 9.4
Corn A 40
6/40 58.068.5 4.5
Waxy rice A 37
6/40/6/40 58.269.0 5.9
Potato A 30
Potato 6/6 35.278.2 13.3
Cassava C 37
6/30 48.878.3 11.7
True yam (Dioscorea) B 32
6/30/6/30 49.778.1 13.1
Taro A 31
6/40 59.278.3 9.9
Sweet potato C 38
6/40/6/40 60.779.5 10.6

To Tc transition temperature range; DH enthalpy; AMP amylopectin.
a
Each number indicates the temperature for one day of storage; 6/40 means one day at
60  C, followed by one day at 40  C.
Source: Silverio J, Fredriksson H, Andersson R, Eliasson A-C, and Aman P (2000) The
effect of temperature cycling on the amylopectin retrogradation of starches with different
amylopectin unit-chain length distribution. Carbohydrate Polymers 42: 175184.
sample size and inability to determine retrogradation in dilute
starch paste. More often DSC coupled with X-ray analysis pro-
vides better performance in characterizing retrogradation.
X-ray analysis
X-ray analysis can be applied to detect the presence and nature
of crystallinity in native starch granules as well as crystals
formed in aging starch gels. In native starch, the crystallinity
is due to the amylopectin components and the crystal domains
are constructed mainly of A chains and outer B chains of
amylopectin. Native starches can be categorized into three
groups according to their X-ray diffraction pattern. The A type
Sources: Zobel (1988b) and Gunaratne A and Hoover R (2002) Effect of heat-moisture
treatment on the structure and physicochemical properties of tuber and root starches.
Carbohydrate Polymers 49: 425437.
crystallinity is found mainly in cereal starches. Most tuber and
root starches exhibit typical B type X-ray pattern and C type is
intermediate between A and B types, shown in legumes, cas-
sava, and some varieties of sweet potato (Table 8). X-ray anal-
ysis of retrograded starch has shown that aging gels form B-type
crystals irrespective of the native starch. Some research using
wide range and small angle X-ray diffraction, shows that crys-
talline formation in starch paste occurred primarily due to
amylopectin aggregation, whereas amylose provides a template
effect and highly ordered amylose aggregation does not neces-
sarily possess a crystalline nature but accelerates the amylopec-
tin reassociation. Recently, more information on bread staling
has been revealed by X-ray analysis. Aggregation of starch poly-
mers that formed crystals in bread staling was shown from
X-ray analysis; however, NMR and FTIR techniques are better
206 CARBOHYDRATES | Starch: Analysis of Quality
Table 9 Gel textural properties of maize starches differing in amylose content
Sample Probe type Hardness (g)
Regular 5 (mm) 75
Waxy
High amylose 19
Waxy 20 (mm) 28
High amylose 128
Starch paste after RVA analysis kept at 25  C for 24 h
Source: Liu H, Ramsden L, and Corke H (1999) Physical properties and enzymatic digestibility of hydroxypropylated ae, wx, and normal maize starch. Carbohydrate Polymers 40:
175182.
at detecting minor differences of starch polymer aggregation
than X-ray analysis.
Rheological methods
Measurement of pasting parameters, analysis of textural proper-
ties of starch paste using texture analyzer, and small deforma-
tion dynamic techniques oscillatory rheometry can be applied
to the behavior of starch. Pasting behavior of starch indicates the
trend of starch retrogradation. In a typical pasting profile, the
magnitude of setback reflects the tendency of mainly amylose
polymer reassociation in a starch paste. Several investigations
have been performed to study the properties of aging breads
using viscometry and have observed a tendency of decreased
peak viscosity when bread staling progresses. Recently there is a
growing interest, especially in industry, in use of texture ana-
lyzers that can directly and rapidly measure the textural proper-
ties such as hardness, stickiness, cohesiveness, adhesiveness,
fracturability of starch paste (Table 9). Long-term reliability
and accuracy, direct measurement, time saving, and technical
feasibility of the texture analyzer increases its attractiveness
among food technologists. The main issue which requires con-
tinuing work is that of instrumental to sensory correspondence,
i.e., whether the texture analysis profile is adequately predictive
of the sensory traits important to the consumer.
Many additional methods can be employed to detect retro-
gradation. No individual method is able to fully characterize
all the events of the starch-aging process. DSC measures only
the transition of heat energy in the melting of crystal aggre-
gates, X-ray measures the presence and characteristics of retro-
graded starch crystals, and texture analysis provides textural
parameters for the aged starch gel. Therefore, it is clear that a
multiple analytical technique approach for the advance detec-
tion of retrogradation behavior is necessary. For example, a
study of bread staling with multiple techniques, DSC, FTIR,
and NIR, led to greater understanding of the process than
would be possible with each individual technique used alone.
Exercises for Revision
What are the chemical and functional differences between
amylose and amylopectin?
Explain the terms waxy and non-waxy in terms of starch
composition and function.
Describe the range of methods available to determine amy-
lose:amylopectin ratio. What situations might suit the use
of one of these methods versus another?
Adhesiveness Springiness Cohesiveness
219 0.94 0.51
18 0.61 0.031
18 0.94 0.81
160 0.85 0.35
Describe the process of starch pasting and the methods for
its analysis. How does this property affect the uses of starch
in the food industry?
What is meant by starch retrogradation? Explain the result-
ing beneficial and undesirable consequences for food
products.
What are the events of the starch-aging process? How do
these phenomena affect the functional use of starch in food
and non-food uses?
Exercises for Readers to Explore the Topic Further
Explore the range of mutant types of grain species for which
wide differences in amylose:amylopectin ratio have been
developed. What are the resulting differences in functional
properties and thus in applications in utilization?
Starch is insoluble in water. Do you agree with this state-
ment? Under what circumstances and for which forms of
starch is the statement true and untrue? Explain how the
solubility (or not) of starch is critical to its utilization.
Explore how the presence of starch-degrading enzymes
(such as amylase enzymes) increases starch solubility and
thus its functional properties.
What lessons might be learned from research into the chain
lengths and degree of branching of the various starch
molecules.
See also: Carbohydrates: Carbohydrate Metabolism; Glycemic
Index, Glycemic Load, and Their Health Benefits; Resistant Starch and
Health; Starch: Chemistry; Starch: Starch Architecture and Structure;
Starch: Synthesis; Food-quality Testing: Noodles: Testing for
Quality; Pasta: Quality Testing Methods; Rice: Eating Quality;
Genetics of Grains: Maize: Other Maize Mutants; Maize: Quality
Protein Maize; Grain Composition and Analysis: Standardized Test
Methods for Grains and Grain-Based Products; The Composition of
Food Grains and Grain-Based Products; Processing of Grains:
Starch: Modification; Starch: Uses of Native Starch; Wheat
Processing: Baked Product Staling: Mechanisms, Determinations,
and Anti-staling Strategies; The Basics: Grain and Plant Morphology
of Cereals and how characters can be used to identify varieties.
Further Reading
Collado LS and Corke H (2003) Starch properties and functionalities. In: Kaletunc G and
Breslauer KJ (eds.) Characterization of Cereals and Flours, pp. 473506. New York:
Marcel Dekker.

Daniel JR and Whistler RL (1985) Principal changes in starch during food processing.
In: Richardson TR and Finley JW (eds.) Chemical Changes in Food During
Processing, pp. 305326. Westport, CT: AVI Publishing Co.
Eliasson A-C (2003) Utilization of thermal properties for understanding baking and
salting processes. In: Kaletunc G and Breslauer KJ (eds.) Characterization of Cereals
and Flours, pp. 65115. New York: Marcel Dekker.
Fredriksson H, Silverio J, Andersson R, Eliasson A-C, and Aman P (1998) The influence
of amylose and amylopectin characteristics on gelatinization and retrogradation
properties of different starches. Carbohydrate Polymers 35: 119134.
Gonera A and Cornillon P (2002) Gelatinization of starch/gum/sugar systems studied by
using DSC, NMR, and CSLM. Starch/Starke 54: 508516.
Gunaratne A and Hoover R (2002) Effect of heat-moisture treatment on the structure and
physicochemical properties of tuber and root starches. Carbohydrate Polymers
49: 425437.
Hoover R, Vasanthan T, Senanayake NJ, and Martin AM (1994) The effects of defatting
and heat-moisture treatment on the retrogradation of starch gels from wheat, oat,
potato, and lentil. Carbohydrate Research 261: 1324.
Jane J, Kasemsuwan T, Leas S, Zobel HF, and Robyt JF (1994) Anthology of starch
granules morphology by scanning electron microscope. Starch/Starke 46: 121.
Karim AA, Norziah MH, and Seow CC (2000) Methods for the study of starch
retrogradation. Food Chemistry 71: 936.
Liu H, Ramsden L, and Corke H (1999) Physical properties and enzymatic digestibility
of hydroxypropylated ae, wx, and normal maize starch. Carbohydrate Polymers
40: 175182.
Perez EE, Breene WM, and Bahnassey YA (1998) Gelatinization profiles of cassava,
sagu, and arrowroot native starches as measured with different thermal and
mechanical methods. Starch/Starke 50: 1416.
CARBOHYDRATES | Starch: Analysis of Quality 207
Phillips DL, Xing J, Liu H, Pan DH, and Corke H (1999) Potential use of Raman
spectroscopy for the determination of amylose content in maize starch. Cereal
Chemistry 76: 821823.
Planchot V, Gerard C, Bertoft E, and Colonna P (2001) An approach to structural
analysis of granules using genetically modified starches. In: Barsby TL, Donald AM,
and Frazier PJ (eds.) Advances in Structure and Function, pp. 104128. UK: The
Royal Society of Chemistry.
Praznik W, Huber A, Watzinger S, and Beck RHF (1994) Molecular characteristics of
high amylose starches. Starch/Starke 46: 8894.
Silverio J, Fredriksson H, Andersson R, Eliasson A-C, and Aman P (2000) The effect of
temperature cycling on the amylopectin retrogradation of starches with different
amylopectin unit-chain length distribution. Carbohydrate Polymers 42: 175184.
Tester RF and Morrison WR (1990) Swelling and gelatinization of cereal starches 1.
Effects of amylopectin, amylose and lipids. Cereal Chemistry 67: 551557.
Yu L and Christie G (2001) Measurement of starch thermal transitions using differential
scanning calorimetry. Carbohydrate Polymers 46: 179184.
Zobel HF (1988) Molecules to granules: a comprehensive starch review. Starch/Starke
40: 4450.
Relevant Website
starch.dk http://www.starch.dk Website of the International Starch Institute, Denmark,
with useful links to other starch-related sites.
 Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
GB Fincher, Australian Research Council Centre of Excellence in Plant Cell Walls, University of Adelaide, Glen Osmond, SA, Australia
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Structure and function of cell wall polysaccharides.
Fine structure of wall polysaccharides and how these vary in
different tissues and at different stages of development.
How fine structure affects physicochemical properties,
including viscosity, solution behavior, and gelation.
Genes involved in the biosynthesis, remodeling, and deg-
radation of cell wall polysaccharides.
The importance of solubility and viscosity in cereal proces-
sing and human health.
The benefits of dietary fiber from cereal grains in reducing
the risk of serious human diseases, including type II diabe-
tes, colorectal cancer, and cardiovascular disease.
Learning Objectives
To know how chemical structure affects solution properties.
To understand how cell wall polysaccharides contribute to
dietary fiber.
To know the structurefunction relationships in cereal pro-
cessing and nutrition.
Introduction
The nonstarchy polysaccharides of cereal grains include cellu-
lose, (1 ! 3, 1 ! 4)-b-D-glucans, heteroxylans (arabinoxylans),
glucomannans, xyloglucans, pectic polysaccharides, callose,
fructans, and arabinogalactan proteins (AGPs). With the excep-
tion of the fructans and the AGPs, all are key components of
the walls of cells in the various tissues that comprise the grain.
In this article, the chemistry and physicochemical properties of
nonstarchy polysaccharides from cereal grains, which are
important in human and animal nutrition and which have
an impact in grain utilization and in cereal technology, will
be described.
The cells of grain tissues have diverse functions during
development, during dormancy, and after germination. The
pericarp and seed coat tissues are concerned with the protec-
tion of the seed during development and dormancy. However,
at grain maturity, these tissues are nonliving and consist almost
entirely of cell wall residues. The nucellar tissue between the
seed coat and the aleurone surface is involved in the transfer of
nutrients to the developing grain but collapses at maturity to
leave compressed cell wall remnants in the mature grain. Cells
of the starchy endosperm are nonliving in the mature grain,
but are packed with starch and storage protein and, in most
cereal species, have relatively thin cell walls. In contrast, the
thick-walled, nucleated, aleurone cells are living at grain
maturity and are packed with membrane fragments, protein
bodies, and lipid droplets. The living embryonic tissues of the
grain are at least partly differentiated into functional cell types
208 Encyclopedia of Food Grains, Second Edition
that are eventually found in shoots and roots. At the interface
of the embryo and the starchy endosperm lies the scutellum,
which delivers nutrients to the developing endosperm and
plays a dual role during germination; in the first instance, the
scutellar epithelial layer secretes degradative enzymes into the
proximal region of the starchy endosperm, but later, it trans-
locates digestion products of the endosperm reserves into the
developing embryo.
The cell wall compositions and organization of the cell
types found in the tissues of the grain reflect these different
functions and, in turn, determine the nutritional impact and
behavior of various grain parts in technical operations during
grain utilization and processing. Together, the nonstarchy
polysaccharides usually constitute <10% by weight of the
grain (Table 1) but can nevertheless be key determinants of
grain quality. Although the precise physical relationships
between individual nonstarchy polysaccharides and other
wall components have not been described, it is generally con-
sidered that in the wall, microfibrils of cellulose are embedded
in a matrix phase that consists mainly of noncellulosic poly-
saccharides and smaller amounts of protein. Lignification of
walls in the outer layers of the grain is observed, but little or no
lignification is found in most cells of the endosperm and
embryo. Wall integrity is maintained predominantly through
extensive noncovalent interactions, especially hydrogen bond-
ing, between the matrix phase and the microfibrillar constitu-
ents. In the walls of some grain tissues, covalent associations
between heteroxylans, lignin, and proteins are present. Indeed,
there have been several recent reports of previously unrecog-
nized covalent linkages between noncellulosic polysaccharides
of plant cell walls more generally. The extent of covalent asso-
ciations between components also varies with the wall type
and genotype. The relative ease of extraction of noncellulosic
polysaccharides from the different wall types in cereal grains is
strongly influenced by both noncovalent and covalent interac-
tions. Cell walls have been isolated and analyzed from some
cereal grains, in particular aleurone and starchy endosperm
walls from wheat and barley (Table 1).
Noncellulosic polysaccharides, especially heteroxylans and
(1 ! 3, 1 ! 4)-b-D-glucans, constitute a relatively high propor-
tion of the walls of the aleurone and starchy endosperm and
probably also of the scutellum. Cellulose contents are corre-
spondingly lower (Table 2). The generally low cellulose con-
tent of these walls, together with the fact that the walls of the
endosperm contain no lignin, might be related functionally to
a limited requirement for structural rigidity of walls in central
regions of the grain and to a requirement for rapid depolymer-
ization of wall components following germination of the
grain. There is some indirect information on the composition
of starchy endosperm walls from rice, which are extremely thin
and appear to consist mostly of cellulose and heteroxylans. The
(1 ! 3, 1 ! 4)-b-D-glucan levels appear to be low, and varying
amounts of pectin, xyloglucan, and glucomannan have been
reported in rice endosperm walls.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00090-5
 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 209

Table 1 Composition of cell walls of cereal grains (% by weight)

Component

(1 ! 3, 1 ! 4)-
Cereal Origin of cell wall Cellulose Glucomannan b-D-Glucan Heteroxylan Pectin Lignin Proteina

Wheat (Triticum Aleurone 2 2 29 65 1

aestivum L.) Starchy endosperm 2 2 20 70
Bran (pericarp, seed 29 6 64 8.3 9.2
coat, and aleurone)
Beeswing bran 30 60 12 6
(outer pericarp)
Barley (Hordeum Aleurone 2 2 26 71 6
vulgare L.) Starchy endosperm 2 2 75 20 5
Rice (Oryza sativa L.) Bran 9 3 6 60 8 12 8
Maize (Zea mays L.) Bran (pericarp, seed 23 67
coat, and aleurone)
a
Values for protein may include cellular proteins.

Table 2 Content of (1 ! 3, 1 ! 4)-b-D-glucan and heteroxylan in whole cereal grains (gram per 100 g DW)

(1 ! 3, 1 ! 4)-b-D-Glucan Arabinoxylan

Cereal Total Water-extractable Total Water-extractable

Wheat (Triticum aestivum L.) 0.52.3 0.02 4.09.0 0.30.9

Durum (Triticum durum) 0.50.6 0 ?
Spelt (Triticum aestivum ssp. spelta) 0.57 ?
Barley (Hordeum vulgare L.) 210 Up to 50% 48 0.4
Hull-less 3.95.4 1.02.7
Waxy hull-less 615
Rice (Oryza sativa L.) 0.13 0.03 ?
Sorghum (Sorghum bicolor L.) 1.16.2 ?
Rye (Secale cereale L.) 1.02.0 7.112.2 0.62.4
Rye-derived wheats (1B/1R translocations) 5.25.5
Maize (Zea mays L.) 0.81.7 5.16.8 4.6
Oats (Avena sativa L.) 3.86.1 3.75.0 2.24.1 0.2
Triticale 0.341.8 0 5.87.4

?, data unavailable.

In walls of cells in the multiple tissue layers of the mater-
nally derived, pericarpseed coat, which provides protection
for the embryo and endosperm and which is not mobilized
during germination, cellulose and lignin contents are much
higher and the concentrations of noncellulosic polysaccharides
are correspondingly lower.
Cellulose
Cellulose is a component of all cell walls in cereal grains.
However, its concentration varies widely. Cellulose is usually
a minor constituent of the primary walls of starchy endosperm
and aleurone cells of cereal grains, but its content is much
higher in secondary walls of the pericarp and seed coat cells
(Table 1). It is a linear homopolymer composed of
b-D-glucopyranosyl residues, all of which are (1 ! 4)-linked
(Figure 1(a)). Individual molecules have a degree of
polymerization of up to 6000 in primary walls and 14 000 in
secondary walls. The molecular chains have an extended,
ribbonlike conformation that allows parallel packing of chains
into three-dimensional, fibrillar aggregates stabilized by exten-
sive intermolecular hydrogen bonding and van der Waals
interactions (Figure 1(b)). These aggregates, termed microfi-
brils, are easily visible under the electron microscope and may
reach diameters of 3 nm in primary walls and 510 nm in
secondary walls. Within microfibrils, individual cellulose mol-
ecules are mostly packed into parallel, highly ordered, crystal-
line arrays, although there are usually regions where molecular
alignment is less ordered. In particular, the surface molecules
may have a different conformational and hydrogen-bonding
arrangement compared with molecules within the more crys-
talline interior. Long-held views that the cellulosic microfibrils
of higher plant walls contain about 36 individual cellulose
molecules have recently been challenged, and it appears likely
that the number of individual cellulosic chains per microfibril
is considerably lower than originally believed.
210 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides

Figure 1 Cellulose structure. (a) Portion of a cellulose molecule. (b) Crystal structure of cellulose I: (i) projection down the fiber axis c showing the
layers (sheets) of cellulose chains hydrogen-bonded in the ac plane but lacking intersheet bonding. The intersheet bonding involves van der Waals
interactions between the hydrophobic faces of the glucose units. (ii) View of a layer approximately perpendicular to the ac plane showing the two
intramolecular hydrogen bonds in the direction of the fiber axis c and the interchain hydrogen bonds. Reproduced from Kroon-Batenburg LMJ and Kroon
J (1995) The crystal and molecular structures of cellulose. Carbohydrates in Europe 12: 1519, with permission.

Cellulose microfibrils may also associate with water and
matrix polysaccharides such as the (1 ! 3, 1 ! 4)-b-D-glucans,
heteroxylans (arabinoxylans), and glucomannans. Cellulose is
insoluble in water but swells in concentrated sodium hydroxide
solutions and may be brought into solution by powerful
hydrogen bond-breaking reagents such as N-methylmorpholine
N-oxide. Thermal and shear treatments, followed by alkaline
peroxidation and shearing to remove lignin, have been used to
convert cellulose-rich maize bran (Table 1) to a cellulosic gel for
use as a dietary fiber supplement.
(1 ! 3, 1 ! 4)-b-D-Glucans
(1! 3, 1 ! 4)-b-D-Glucans, also referred to as mixed-linkage or
cereal b-glucans, occur almost exclusively in members of the
monocotyledon family Poaceae, to which the cereals and grasses
belong, and in related families of the order Poales (sensu stricto).
They are important constituents of the walls of the starchy
endosperm and aleurone cells of many cereal grains, where
they can account for up to 70% by weight of the walls (Table 1).
The concentration of (1! 3, 1 ! 4)-b-D-glucans in cereal grains
varies across the grain and depends on the genotype, the posi-
tion of the grain on the spike and environmental factors such as
planting location, climatic conditions during grain develop-
ment, and soil nitrogen. Barley, oat, and rye grains are rich
sources of (1! 3, 1 ! 4)-b-D-glucans, whereas wheat, rice, and
maize have lower concentrations of the polysaccharide
(Table 2). The (1! 3, 1 ! 4)-b-D-glucans are relatively minor
components of walls in vegetative tissues of cereals and grasses.
A structurally related (1! 3, 1 ! 4)-b-D-glucan, lichenin, is
found in the walls of the fungal component of the lichen,
Iceland moss (Cetraria islandica). There have been recent reports
of the presence of (1! 3, 1 ! 4)-b-D-glucans in the walls of the
horsetail fern (Equisetum spp.), in some bryophytes, various
algal species, and the walls of certain fungal pathogens.
Structure
(1 ! 3, 1 ! 4)-b-D-Glucans are linear, unbranched polysaccha-
rides in which b-D-glucopyranosyl monomers are polymerized
through both (1 ! 4)- and (1 ! 3)-linkages (Figure 2(a)). The
ratio of (1 ! 4) to (1 ! 3)-linkages is generally fairly constant,
and in the range 2.22.6:1, although in the (1 ! 3, 1 ! 4)-b-D-
glucan from sorghum endosperm, the ratio is 1.15:1. The two
types of linkages are not arranged in regular, repeating
sequences. Single (1 ! 3)-linkages are usually separated by
two or more (1 ! 4)-linkages. Regions of two or three adjacent
(1 ! 4)-linkages predominate, but there is no regularity in the
arrangement of these units. Thus, cereal (1 ! 3, 1 ! 4)-b-D-
glucans may be considered as (1 ! 3)-b-linked copolymers of
cellotriosyl (G4G4GRed), cellotetraosyl (G4G4G4GRed) units,
and longer (1 ! 4)-b-D-oligoglucosyl units (Figure 2(b)). The
ratio of tri- to tetrasaccharide units varies between cereal spe-
cies, as follows: wheat (3.04.5:1), barley (2.93.4:1), rye
(2.7:1), and oats (1.82.3:1). The ratios vary according to
the temperature and conditions of extraction. However,
up to 10% of the chain may consist of longer stretches of
520 adjacent (1 ! 4)-linkages, and at low frequency, single
(1 ! 4)-linkages are detected. An important point related to
(1 ! 3, 1 ! 4)-b-D-glucan structure is that the single (1 ! 3)-
b-D-glucosyl residues will introduce molecular kinks into the
main chain of the polysaccharide and thereby interrupt what
would otherwise be an extended, straight cellulose molecule
 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 211

Figure 2 (a) Portion of a (1 ! 3, 1 ! 4)-b-D-glucan molecule. (b) Distribution of linkages in a (1 ! 3, 1 ! 4)-b-D-glucan. (G, b-glucosyl unit);
3 (1 ! 3)-linkage, 4 (1 ! 4)-linkage; red, reducing end, arrows, site of hydrolysis by (1 ! 3, 1 ! 4)-b-D-glucan endohydrolase (EC 3.2.1.73).
(c) Perspective drawings of computer-generated, instantaneous conformations of b-glucans: (i) (1 ! 4)-b-D-glucan (cellulose), (ii) (1 ! 3)-
b-D-glucan (callose), (iii) (1 ! 3, 1 ! 4)-b-D-glucan, () (1 ! 4)-linked residues, and () (1 ! 3)-linked residue. Reproduced from Fincher GB
and Stone BA (1986) Cell walls and their components in cereal grain technology. Advances in Cereal Science and Technology 8: 207295, with
permission.

(Figure 2(c) (i) cf. Figure 2(c) (iii)). First-order Markov chain
analyses have been used to show that the DP3 and DP4 units
of the water-soluble barley (1 ! 3, 1 ! 4)-b-D-glucan are
arranged at random (Figure 3), which means that the molec-
ular kinks imposed by (1 ! 3)-b-D-glucosyl residues in the
main chain of the polysaccharide will also be irregularly
spaced.
Solubility
The extractability of (1 ! 3, 1 ! 4)-b-D-glucans from walls of
cereal grains is partly a function of their degree of self-
association and their association with other wall polysaccha-
rides and proteins. In addition, extractability depends on the
molecular mass and linkage distribution in the (1 ! 3, 1 ! 4)-
b-D-glucan chains. Extensive association with other polymers
and very high molecular masses will render the (1 ! 3, 1 ! 4)-
b-D-glucans more difficult to extract from the grain.
The average molecular masses reported for cereal (1 ! 3,
1 ! 4)-b-D-glucans range from 48 000 (degree of polymeriza-
tion, DP,  300) to 3 000 000 (DP  1850), depending on the
cereal species, cell wall type, extraction procedure, and the
method used for molecular mass determination. It is likely
that in many cases, the DP of the polysaccharide has been over-
estimated, particularly where gel permeation chromatography is
used for size determination, because standards used to calibrate
the columns often have a more globular conformation than the
..G 3 G 4 G 4 G 3 G 4 G 4 G 4 G 3 G 4 G 4 G 3 G ..
1 p
p Cellotriosyl Cellotetraosyl 1 q
q
r=p+q
r = autocorrelation (interdependence between
DP3 and DP4 units)
Figure 3 Determination of the arrangement of cellotriosyl (DP3) and
cellotetraosyl (DP4) residues in a water-soluble (1 ! 3, 1 ! 4)-
b-D-glucan from barley grain. The polysaccharide was hydrolyzed
(vertical arrows) with a specific (1 ! 3, 1 ! 4)-b-D-glucan
endohydrolase until the penultimate oligosaccharides of DP6, DP7, and
DP8 predominated. The autocorrelation coefficient (r) is defined in
terms of the probabilities that a cellotriosyl residue is followed by
another cellotriosyl residue (p) or a cellotetraosyl residue (q), which
can be estimated from the abundance of the oligosaccharide products.
If the polysaccharide is treated as a first-order Markov chain, a r value
of 1 indicates an adjacent arrangement of DP3 and DP4 units, a r
value of 1 indicates an alternating arrangement of DP3 and DP4 units,
and a r value of 0 indicates a random arrangement of DP3 and DP4
units. Values very close to 0 were obtained for two barley (1 ! 3,
1 ! 4)-b-D-glucans.
212 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
highly asymmetrical wall polysaccharides. The (1 ! 3, 1 ! 4)-b-
D-glucans are invariably polydisperse with respect to molecular
mass, and this is illustrated by a weight average to number
average molecular mass ratio (Mw/Mn) of 1.18 for barley
(1! 3, 1 ! 4)-b-D-glucan. Certain barley (1 ! 3, 1 ! 4)-b-D-
glucans are covalently associated with small amounts of protein
and have estimated molecular masses of up to 40 000 000.
Apart from their molecular mass, the fine structure of the
(1 ! 3, 1 ! 4)-b-D-glucans is a key determinant of their solu-
bility. There are two components of fine structure that need to
be considered in this context. Firstly, the longer stretches of
520 adjacent (1 ! 4)-linkages in the polysaccharide will be
cellulosic in nature and would allow intermolecular aggrega-
tion and hence reduced solubility. However, these longer
blocks of adjacent (1 ! 4)-linkages are not abundant and
their effects on solubility have not been quantitated. The sec-
ond and probably more important element of polysaccharide
fine structure that will affect solubility is the relative propor-
tions of the major subunits of the (1 ! 3, 1 ! 4)-b-D-glucans,
namely, the cellotriosyl (DP3) and cellotetraosyl (DP4) units
(Figure 2(b)). The ratio of these tri- to tetrasaccharide units
(DP3:DP4 ratio) as illustrated in Figure 2(b) is proving to be a
useful predictor of (1 ! 3, 1 ! 4)-b-D-glucan solubility, insofar
as the polysaccharide appears to become less soluble in aque-
ous media as the ratios move away from 1:1. As the number of
DP3 units in the (1 ! 3, 1 ! 4)-b-D-glucan increases, the
molecular kinks imposed by the single (1 ! 3)-b-D-glucosyl
residues will become more evenly spaced, and the polysaccha-
ride chains will be able to align over longer regions. Thus,
solubility will decrease as the DP3:DP4 ratio increases to larger
values. Similarly, as the number of DP4 units in the (1 ! 3,
1 ! 4)-b-D-glucan increases, the molecular kinks imposed by
the single (1 ! 3)-b-D-glucosyl residues will again become
more evenly spaced, and solubility will decrease as the DP3:
DP4 ratio decreases to low values. This key argument for
explaining (1 ! 3, 1 ! 4)-b-D-glucan solubility in terms of
DP3:DP4 ratios only holds if the major DP3 and DP4 units
of the polysaccharide are arranged in an essentially random
fashion along the (1 ! 3, 1 ! 4)-b-D-glucan chain. This has
been demonstrated for very few (1 ! 3, 1 ! 4)-b-D-glucans,
but a random arrangement of the DP3 and DP4 units does
occur in a water-soluble (1 ! 3, 1 ! 4)-b-D-glucan from barley
grain (Figure 3).
The variations in the DP3:DP4 ratios between (1 ! 3,
1 ! 4)-b-D-glucans from different tissues and at different devel-
opment stages indicate that the biosynthetic enzymes respon-
sible for (1 ! 3, 1 ! 4)-b-D-glucan synthesis are able to adjust
the fine structure and hence the physicochemical properties of
the polysaccharide. There is circumstantial evidence that the
amount and solubility of (1 ! 3, 1 ! 4)-b-D-glucan in grain are
correlated (Table 3). However, it needs to be emphasized that
the solubility of the (1 ! 3, 1 ! 4)-b-D-glucan will also depend
on intermolecular interactions with other (1 ! 3, 1 ! 4)-b-D-
glucan molecules and with other polysaccharides and proteins
in the grain during cereal processing, as noted earlier.
A portion of the (1! 3, 1 ! 4)-b-D-glucan from barley, oat,
and rye flours is extracted by water at pH 7.0 and 40  C. These
flours consist predominantly of aggregates from starchy
endosperm cells. Further fractions can be solubilized at higher
temperatures. The proportion of total (1 ! 3, 1 ! 4)-b-D-glucan
Table 3 Adoption of (1 ! 3, 1 ! 4)-b-D-glucan in grain is correlated
with fine structure
Typical properties of
Species (1,3;1,4)-b-glucans in grain
Oats (Avena sativa) Ratio in grain 2:1
80% soluble in water (40  C)
68% (w/w) grain
Barley (Hordeum vulgare) Ratio in grain 2.8:1
20% soluble in water (40  C)
410% (w/w) grain
Wheat (Triticum aestivum) Ratio in grain 3.2:1
0% solubility in water (40  C)
0.52.3% (w/w) grain
As the DP3:DP4 ratio increases, the solubility of the (1 ! 3, 1 ! 4)-b-D-glucan
decreases and the amount in the grain also appears to decrease.
that is water-soluble at 40  C varies within and between species.
For example, waxy (high-amylose) barleys have a higher pro-
portion of water-soluble (1 ! 3, 1 ! 4)-b-D-glucan than normal
barleys. (1! 3, 1 ! 4)-b-D-Glucans extracted from barley at
40  C have a slightly lower tri- or tetrasaccharide ratio (1.7:1)
than those extracted at 65  C (2.0:1). Complete extraction of
cereal (1! 3, 1 ! 4)-b-D-glucans from grain requires the use of
alkaline extractants such as 4 M NaOH or aqueous Ba(OH)2,
containing NaBH4 to prevent alkali-induced degradation from
the reducing terminus. Alkali-extracted barley (1 ! 3, 1 ! 4)-
b-D-glucan fractions have higher molecular masses, higher ratios
of (1! 4):(1 ! 3)-linkages, more contiguously linked (1 ! 4)-
linked segments, and higher trisaccharidetetrasaccharide ratios
than their water-extractable counterparts. Other extractants,
such as dimethyl sulfoxide, hot perchloric acid, trichloroacetic
acid, N-methylmorpholine N-oxide, and dimethylacetamide
LiCl, have also been used to solubilize (1 ! 3, 1 ! 4)-
b-D-glucans, but all are liable to cause some depolymerization
or degradation of the polymer. Thus, care must be exercised in
comparing structural differences in cereal (1 ! 3, 1 ! 4)-
b-D-glucans extracted using different solvents. Once extracted
with hot water or alkali, the (1 ! 3, 1 ! 4)-b-D-glucans are
soluble at neutral pH and room temperature. However, upon
cooling, the (1! 3, 1 ! 4)-b-D-glucans slowly aggregate and
precipitate.
Molecular Conformations
In aqueous media barley (1 ! 3, 1 ! 4)-b-D-glucans adopt a
partially extended, somewhat rigid, reptate (snakelike) chain
conformation (Figure 2(c)) with a cross-sectional diameter of
0.45 nm and a lengthwidth (axial) ratio of  100. It is likely
that other (1 ! 3, 1 ! 4)-b-D-glucans adopt a similar confor-
mation in aqueous media, but there are insufficient conforma-
tional data available for (1 ! 3, 1 ! 4)-b-D-glucans from a
range of sources to allow meaningful comparisons. It would
be particularly interesting to compare the molecular confor-
mations of (1 ! 3, 1 ! 4)-b-D-glucans with different DP3:DP4
ratios.
In the solid state, x-ray fiber diagrams of the water-soluble
barley, (1 ! 3, 1 ! 4)-b-D-glucan can be interpreted as a
(1 ! 3)-linked cellotriosyl polymer with a threefold screw
 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
axis and a fiber repeat of 4.2 nm. The shape of the chain is
reminiscent of cellulose in the cellotriosyl and longer cello-
oligosaccharide units, but the (1 ! 3)-linkages provide some
helical character.
Viscosity of Solutions of (1 ! 3, 1 ! 4)-b-D-Glucans
The intrinsic viscosities of (1 ! 3, 1 ! 4)-b-D-glucan solutions
are dependent on the cereal species from which the polysac-
charide is extracted and on the solvent used. Intrinsic viscosity
values for barley (1 ! 3, 1 ! 4)-b-D-glucans in water range
from 4.6 to 6.9 dl g1 and from 2.0 to 9.6 dl g1 for oat
(1 ! 3, 1 ! 4)-b-D-glucans. The extractability, solubility, and
viscosity characteristics of cereal (1 ! 3, 1 ! 4)-b-D-glucans
can be understood by reference to their fine structures, as
noted earlier. Long runs of adjacent (1 ! 4)-b-D-glucosyl
units in the (1 ! 3, 1 ! 4)-b-D-glucan chain will form extended
ribbonlike cellulosic stretches, but the insertion of a (1 ! 3)-
linked glucosyl unit between them causes a kink in the mol-
ecule (Figure 2(c)). Chain flexibility arises principally from the
isolated (1 ! 3)-b-linkages. Because the (1 ! 3)-linked
b-glucosyl units do not occur regularly along the chain, the
polysaccharides are not able to align over extended regions and
hence remain dispersed and soluble in aqueous media. The
overall extended polysaccharide conformation is due to the
predominance of (1 ! 4)-linkages and results in the occupancy
of a high volume of solvent (high hydrodynamic volume). This
results in aqueous solutions of high viscosity.
The viscosity of solutions of cereal (1 ! 3, 1 ! 4)-b-D-
glucans depends strongly on concentration. At concentrations
of oat (1 ! 3, 1 ! 4)-b-D-glucan where the total volume occu-
pied by the chains is, in aggregate, less than the total volume of
the solution, there is contact between individual chains, but
the viscosity of the solution is independent of shear rate. At
concentrations above the critical overlapping concentration,
there is interpenetration of the chains in response to increasing
space occupancy. The viscosities of such aqueous solutions are
reduced with increasing shear rate, a phenomenon known as
shear thinning. These different concentration-dependent rhe-
ological properties of (1 ! 3, 1 ! 4)-b-D-glucan solutions may
be of significance in their physiological responses as food
ingredients and in their use in food formulations.
Gelation
(1 ! 3, 1 ! 4)-b-D-Glucan solutions at concentrations of 5%
(w/v) or more form elastic gel networks. The gels are thermo-
reversible, exhibit broad melting transitions, and show syner-
esis. Their melting temperatures depend on the source of
(1 ! 3, 1 ! 4)-b-D-glucan as follows: lichenin, 73  C; barley,
65  C; and oats, 62  C. These values are again related to
trisaccharidetetrasaccharide ratios, which in these examples
are 22:1, 3:1, and 2:1, respectively. In the gel state, the (1 ! 3,
1 ! 4)-b-D-glucan chains interact to form three-dimensional
networks. Association between chains at junction zones,
through pairs of consecutive cellotriosyl units, is proposed.
This accounts for the higher melting temperature of lichenin
gels, where cellotriosyl units make up most of the molecule. In
the cereal (1 ! 3, 1 ! 4)-b-D-glucans, consecutive cellotriosyl
units are less frequent, although the longer runs of adjacent
213
(1 ! 4)-b-D-glucosyl units, not present in lichenin, may also
participate in junction zone formation. The induction time for
gelation depends on mobility and diffusivity. It increases with
the molecular mass of (1 ! 3, 1 ! 4)-b-D-glucan and decreases
as the concentration increases. Gelation is repressed at high
and low temperatures, where molecular motion is high and
low, respectively. Gelation may be induced by homogeniza-
tion, centrifugation, and freezing and thawing.
StructureFunction Relationships in Cereal Processing
and Nutrition
The viscosity-generating properties of soluble cereal (1 ! 3,
1 ! 4)-b-D-glucans are critical determinants in many aspects
of cereal processing. For example, incompletely degraded
(1 ! 3, 1 ! 4)-b-D-glucans from malted barley and cereal
adjuncts can contribute to wort and beer viscosity and are
associated with problems in wort separation and beer filtra-
tion. In other applications, (1 ! 3, 1 ! 4)-b-D-glucans can be
cast as biodegradable films, have foam- and emulsion-
stabilizing ability, and, when used as a flour supplement,
increase water absorption in doughs. Soluble cereal (1 ! 3,
1 ! 4)-b-D-glucans have antinutritive effects in monogastric
animals such as pigs and poultry. The antinutritive effects
have been attributed to the increased viscosity of gut contents,
which is believed to slow both the diffusion of digestive
enzymes and the absorption of degradative products of
enzyme action. This, in turn, leads to slower growth rates.
Moreover, in dietary formulations for poultry, high (1 ! 3,
1 ! 4)-b-D-glucan concentrations are associated with sticky
feces, which are indicative of the poor digestibility of the
(1 ! 3, 1 ! 4)-b-D-glucans and which may present major han-
dling and hygiene problems for producers.
Cereal (1 ! 3, 1 ! 4)-b-D-glucans are important compo-
nents of dietary fiber in human and animal diets. Humans
and monogastric animals produce no enzymes that degrade
(1 ! 3, 1 ! 4)-b-D-glucans, although there are indications that
some depolymerization occurs in the stomach and small
intestine, presumably due to the activity of commensal
microorganisms. By comparison, the soluble (1 ! 3, 1 ! 4)-
b-D-glucans and other nonstarchy polysaccharides are readily
fermented by colonic microorganisms and make a small con-
tribution to digestible energy. In contrast to their antinutritive
effects in monogastric animals, oat and barley (1 ! 3, 1 ! 4)-
b-D-glucans at high concentrations in human foods have
beneficial effects, especially for noninsulin-dependent dia-
betics, by flattening glucose and insulin responses that follow
a meal. High concentrations of (1 ! 3, 1 ! 4)-b-D-glucans
(20% w/v) in food have also been implicated in the reduction
of serum cholesterol concentrations, by lowering the uptake of
dietary cholesterol or resorption of bile acids from the intes-
tine. There is now a good deal of information on the micro-
biota of the human large intestine, on how the relative
abundance of constituent microorganisms is controlled by
the carbon source embodied in dietary fiber, and on how the
fermentation products of specific bacteria affect bowel health
and reduce the risk of serious human diseases, including type II
diabetes, cardiovascular disease, and colorectal cancer. More
recently, connections between dietary fiber and protection
against inflammatory diseases are emerging.
214 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
(1 ! 3, 1 ! 4)-b-D-Glucans, in common with a number of
other polysaccharides, in particular (1 ! 3)-b-D-glucans, mod-
ify immunologic responses in humans by a process that is
mediated through binding to receptors on cells of the reticulo-
endothelial system (leucocytes and macrophages). In addition,
they may have the capacity to activate the proteins of the
human complement pathway, a system that is invoked as a
first line of defense before circulating antibodies are produced.
Analysis and Detection
Specific reagents for the analysis and detection of cereal (1 ! 3,
1 ! 4)-b-D-glucans include enzymes, fluorochromes, and
monoclonal antibodies. The (1 ! 3, 1 ! 4)-b-D-glucan endo-
hydrolase (EC 3.2.1.73) from Bacillus subtilis has been adopted
for the specific quantitation of (1 ! 3, 1 ! 4)-b-D-glucan in
grain or grain extracts. This enzyme cleaves only (1 ! 4)-
linkages that adjoin (1 ! 3)-substituted glucosyl units in the
glucan chain and therefore liberates a series of oligoglucosides
with (1 ! 3)-linked reducing units (Figure 2(b)). These may
be quantified by b-glucosidase cleavage and measurement of
the released glucose with the glucose oxidaseperoxidase
reagent, by separation and quantitation by high-performance
liquid chromatography (HPLC), or by matrix-assisted laser
desorption ionization mass spectrometry (MALDI-TOF-MS).
The latter procedures allow an assessment of the profile of
oligosaccharides making up the primary structure of the (1 ! 3,
1 ! 4)-b-D-glucan and, in particular, the trisaccharide
tetrasaccharide ratio.
b-D-Glucans, including (1 ! 4)-b-D-glucan, cellulose,
(1! 3)-b-D-glucan, callose, and cereal (1 ! 3, 1 ! 4)-b-D-glu-
cans, form complexes with the diphenyldiazo dye, Congo red,
and members of the calcofluor family of diaminostilbene sul-
fonates. These reagents complex poorly with heteroxylans and
AGPs. Irradiation of the complexes induces a brilliant red fluo-
rescence in the case of Congo red (emission lmax, 590 nm; green
excitation, 470 nm) and a bright blue fluorescence with
calcofluor (emission lmax 420, 442 nm; blue excitation,
350 nm). The fluorochromes complex with and precipitate sol-
uble b-glucans. Although complexing with the fluorochromes is
not specific for (1! 3, 1 ! 4)-b-D-glucans, it is nevertheless
useful in quantifying soluble (1! 3, 1 ! 4)-b-D-glucans with
molecular weights of > 10 kDa in grain extracts and in products
such as wort, where cellulose will not be present.
The fluorochromes and a (1 ! 3, 1 ! 4)-b-D-glucan-specific
monoclonal antibody can be used to locate (1 ! 3, 1 ! 4)-
b-D-glucans in tissue sections in histological studies. Although
near-infrared reflectance (NIR) spectra of polysaccharides are
complex, making their direct interpretation difficult, principal
component analysis of the spectra of different polysaccharides
enables characteristic NIR wavelengths to be determined. NIR
spectroscopy has been applied to the determination of (1 ! 3,
1 ! 4)-b-D-glucans in ground barley grain and this method
forms the basis of rapid, routine (1 ! 3, 1 ! 4)-b-D-glucan
estimation.
Genetics
(1 ! 3, 1 ! 4)-b-D-Glucan concentrations in grain are influ-
enced by both genotype and environment. Quantitative trait
locus (QTL) mapping has enabled regions of the barley
genome that control (1 ! 3, 1 ! 4)-b-D-glucan concentrations
to be identified and placed on high-density genetic maps.
Furthermore, the conservation of gene order (synteny) across
cereal genomes means that regions controlling (1 ! 3, 1 ! 4)-
b-D-glucan concentrations in cereals other than barley can
often be deduced from the genetic positions of barley QTLs.
Using a comparative genomics approach based on barley QTLs
and the rice genome sequence, it has been shown that (1 ! 3,
1 ! 4)-b-D-glucan synthesis in the cereals is mediated by the
cellulose synthase-like (Csl) genes of the CslF, CslH, and CslJ
clades. The CslF gene subfamily is the largest of these clades.
There are ten CslF genes in barley and similar numbers in
wheat, sorghum, and rice. The CslH and CslJ gene subfamilies
are generally much smaller. The multiple members of the CslF
gene subfamily are distributed across cereal genomes, although
there is a cluster of CslF genes on chromosome 2H of barley
and in syntenic regions of other cereal genomes. The function
of the gene cluster is not known.
Expression analyses indicate that the CslF6 gene is the most
highly transcribed member of the gene subfamily in develop-
ing barley grain and in most other tissues of barley. The CslF6
gene also appears to be important in most other cereals. Over-
expression of the CslF6 gene in transgenic barley led to signif-
icantly elevated levels of (1 ! 3, 1 ! 4)-b-D-glucan in the grain
and to increased dietary fiber values. This work also suggested
that different CslF isoenzymes synthesize (1 ! 3, 1 ! 4)-b-D-
glucans with different fine structures, as measured by DP3:DP4
ratios. Genome-wide association studies are showing that addi-
tional genes are likely to participate in the regulation of (1 ! 3,
1 ! 4)-b-D-glucan levels in grain.
Heteroxylans
The cereal heteroxylans, or pentosans, are of two main types:
the arabinoxylans and the glucuronoarabinoxylans. They are
characteristically abundant in walls of vegetative tissues of
cereals and grasses, where they represent the core non-
cellulosic polysaccharide in walls of the Poaceae. The arabinox-
ylans are the major noncellulosic polysaccharides in walls of
the starchy endosperm cells and of the aleurone layer of cereal
grains, whereas the glucuronoarabinoxylans are characteristi-
cally found in walls of the pericarpseed coat tissues (Table 1).
Within a cereal species, heteroxylan concentration is influ-
enced by both genotypic and environmental factors.
Structure
Cereal heteroxylans have a (1 ! 4)-b-D-xylan backbone that
has the same shape as the (1 ! 4)-b-D-glucan, cellulose
(Figure 1). The b-xylopyranosyl (Xylp) units of the xylan back-
bone are modified by side-chain branching (substitution)
(Figure 4). The major side chains comprise single a-L-
arabinofuranosyl (Araf) units, situated predominantly at
C(O)3, but in certain species also at C(O)2 of the Xylp units.
In some cases, Araf substitutions occur at both C(O)3 and
C(O)2. The frequency of Araf substitution depends on the
cereal species and wall type and is reflected in a very wide
range of Xylp:Araf ratios. The Xylp units of cereal arabinoxylans
 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 215

may be acetylated. In sorghum heteroxylans, acetyl groups are
present on both the C(O)2 and the C(O)3 of xylopyranosyl
residues. Heteroxylans from cell walls in vegetative tissues,
especially lignified walls such as those from parenchyma, scle-
renchyma fibers, and the pericarpseed coat tissues that are
found in cereal brans, have low degrees of Araf substitution
and therefore high Xylp:Araf ratios. On the other hand, hetero-
xylans from the aleurone layer and starchy endosperm have
relatively low Xylp:Araf ratios.
The distribution of Araf units along the xylan chain is not
regular. In one wheat arabinoxylan fraction, three major types
of substitution patterns have been identified in different
regions of the xylan backbone (Figure 5). In region I, isolated
unsubstituted Xylp units are separated by one or two mono- or
disubstituted units. Region II contains high amounts of Araf
units linked at C(O)3 and region III contains high amounts of
contiguous (up to six and possibly more) unsubstituted Xylp
units.
HO
HO O OH
OH O mers are encountered in fiber from wheat, barley, rye, spelt, oats,
O
O
O rice, and millet. Their structures are shown in Figure 6.
O OH O
O
O bridges also cross-link arabinoxylans, and in lignified walls, the
OH
O O CH2
OH
Other monosaccharides found on the xylan backbone
include D-glucuronic acid (GlcAp) and its 4-O-methyl ether,
linked to the C(O)2 of Xylp units. GlcAp units are found on
endosperm and bran heteroxylans of rice and sorghum and on
bran heteroxylans of wheat, rye, and maize. The barley husk
arabinoxylan has 4% by weight GlcAp. Various oligomeric
chains are also found as substituents on arabinoxylans
(Table 4).
Characteristically, a proportion of the Araf units in the arabi-
noxylans of cereals, grasses, and several other families of
monocotyledons are esterified with the hydroxycinnamic acids,
ferulic acid (FA), and, to a lesser extent, its nonmethoxylated
analog p-coumaric acid (pCA) (Figure 4). These hydroxycinna-
mates are found at C(O)5 of Araf units that are linked to C(O)3
of the Xylp units. Hydroxycinnamates constitute 1.8% (w/w) in
walls of wheat aleurone cells and the FA:pCA ratio is 9:1. In
contrast, walls of the starchy endosperm cells contain only
0.04% (w/w) FA, with a trace of pCA. During wall formation,
enzyme-catalyzed, radical (oxidative) coupling of feruloyl units
on neighboring arabinoxylan chains occurs and leads to their
cross-linking. At least five different cross-linking FA dehydrodi-
In walls of pericarp and seed coat tissues, dehydrodiferulate
arabinoxylans are cross-linked to lignin through FA and dehy-
drodiferulate esterether bridges (Figure 7). Thus, the lignified

Table 4 Substituents on xylan backbone of heteroxylans

a-L-Araf-(l-
H3CO 4-Me-D-GlcpA-(l-
OH D-GlcpA-(l-
b-D-Galp-(l ! 5)-L-Araf-(l-
Figure 4 Structure of portion of a (1! 4)-b-D-xylan chain substituted at
b-D-Xylp-(l ! 2)-L-Araf-(l-
O3-L-arabino of one residue with a 5-O-trans-feruloyl-L-arabinofuranosyl
b-D-Xylp-(l ! 3)-L-Araf-(l-
substituent. Reproduced from Fincher GB and Stone BA (1986) Cell walls
b-D-Galp(l ! 4)-b-D-Xyl-(l ! 2)-L-Araf-(l-(or L)
and their components in cereal grain technology. Advances in Cereal
4Me-D-GlcA-(l ! 4)-D-Xylp-(l ! 4)-D-Galp-(l-
Science and Technology 8: 207295, with permission.

Region I ~ ~

>

Region II ~ ~

>

Region III ~ ~

>

Figure 5 Model of distribution of substituents on an arabinoxylan fraction from wheat endosperm showing three regions with different substitution
patterns. Reproduced from Izydorczyk MS and Biliaderis CG (1995) Cereal arabinoxylans Advances in structure and physico-chemical properties.
Carbohydrate Polymers 28: 3348, with permission.
216 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides

O O
Ara
O O O
Ara Ara
O
O O O
Ara Ara O Ara 8
O H 8 O H 5 OMe
5 O 4 H O
5 OMe OMe
OH
OH
OMe OMe
OH OMe
1 (5-5) 2 (8-O-4) OH 3(8-5)

OH
OMe O O
O Ara
O O Ara
Ara O
H 8
8 H
Ara 4 5
O O O OMe
MeO OMe OH
OH
4 (8-8) 5 (4-O-5)

Figure 6 Ferulated arabinoxylan chains in cell walls of grasses become cross-linked by radical coupling of ferulate monomers into ferulate
dehydrodimers (structures 16). Dotted arrows indicate potential sites for further radical coupling with hydroxycinnamyl alcohols or lignin
oligomers, resulting in cross-linking of arabinoxylans to lignin. Ara is an arabinofuranosyl residue on an arabinoxylan. Reproduced from Grabber
JH, Hatfield RD, Ralph J, Zon J, and Amrhein N (1995) Ferulate cross-linking in cell walls isolated from maize cell suspension cultures. Phytochemistry
40: 10771082, with permission.

Direct ether OH

Direct ester linkage (c)

linkage O=C
O Hydroxycinnamic
Lignin Po acid ester
lys
ac
O ch
O ari
(b) de
C OH
aride

O O
(a) (c) C=O
acch

(d)
C=O
Polys

O OH Hydroxycinnamic
O O acid ether
Lignin
=
=

O O C
O C (e)
O
OH O H
C
O Ferulic acid
=

O (f)
(h) bridge
de

Po O=C
hari

lys OH O
ac O
sacc

ch O
=

Dehydrodiferulic ari C C
de O
Poly

acid diester-ether
=

O O
bridge H
(g)
Dehydrodiferulic
acid diester bridge

: p-Coumaroyl : Feruloyl : Dehydrodiferuloyl

Figure 7 Schematic diagram showing possible covalent cross-links between polysaccharides and lignin in walls. () p-Coumaric acid; ( ) ferulic
acid; ( ) dehydrodiferulic acid. (a) Direct ester linkage; (b) direct ether linkage; (c) hydroxycinnamic acid esterified to polysaccharide; (d)
dehydrocinnamic acid esterified to lignin; (e) hydroxycinnamic acid etherified to lignin; (f) ferulic acid esterether bridge; (g) dehydrodiferulic acid
diester bridge; and (h) dehydrodiferulic acid diesterether bridge. Reproduced from liyama K, Lam TB-T, and Stone BA (1994) Covalent cross-links in
cell wall. Plant Physiology 104: 315320, with permission.
 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
walls in the pericarpseed coat layers and husk of cereal grains
contain significant concentrations of ligninheteroxylan com-
plexes. Wheat bran, which consists largely of pericarpseed
coat and aleurone, contains 0.090.18% (w/w) diferulate res-
idues. This arises almost exclusively from the pericarpseed
coat, because the contribution of FA dimers from the aleurone
is very low (0.03% w/w).
The heteroxylans in maize bran contain 1.3% (w/w) dehy-
drodiferulates, which represent  15 dehydroferulate cross brid-
ges per heteroxylan molecule. Ligninheteroxylan associations
through esterether FA and di-FA cross-links also occur in ligni-
fied walls in stem tissues of cereals and grasses. In walls of wheat
aleurone, highly branched arabinoxylan appears to be cova-
lently associated with protein, possibly through ester-linked FA
that is linked to tyrosine in wall proteins by radical coupling.
Heteroxylans in maize and rye brans are also covalently associ-
ated with protein. The covalent bridges between heteroxylans,
wall proteins, and lignins are important determinants of the
cohesiveness of the wall fabric and of the resistance of lignified
walls to digestion by polysaccharide and ester hydrolases.
Molecular Parameters
Average molecular masses of cereal heteroxylans range from
65 000 to 5 000 000 (DP 50038 000) and, as is the case with
the (1 ! 3, 1 ! 4)-b-D-glucans, the value depends on the cereal
species, cell wall type, extraction procedure, and the method
used for molecular mass determination. The cereal heteroxy-
lans are highly polydisperse with respect to molecular mass as
indicated by Mw/Mn values that range from 1.3 to 4.2 for alkali-
soluble wheat arabinoxylan, 4.1 for water-extractable wheat
arabinoxylan, and 8.5 for rye arabinoxylan.
As with the (1 ! 3, 1 ! 4)-b-D-glucans, a portion of the
arabinoxylans from walls of cereal grains is soluble in water
at 40  C and higher temperatures, but other extractants are
required to bring all the arabinoxylan into solution. Whereas
0.05 MNa2CO3, 8 M urea, and dimethyl sulfoxide extract only
a small part of the water-insoluble arabinoxylan, most can be
extracted with 1 M NaOH, 1 M hydroxylamine hydrochloride,
saturated Ba(OH)2, or 4-methylmorpholine N-oxide. Of these,
saturated Ba(OH)2 containing NaBH4 is the most successful
extractant. Oxidative degradation with chlorite or alkaline
H2O2 is effective in extracting arabinoxylan cross-linked to
lignin from cereal brans. Arabinoxylans extracted under alka-
line conditions may lose all or part of their acetyl and hydro-
xycinnamate substituents.
Solid-State and Solution Conformations
Unsubstituted (1 ! 4)-b-D-xylans can adopt a twofold, helical
conformation with only one hydrogen bond (O50 H-O30 )
between adjacent Xylp units. This conformation is more flexi-
ble than the twofold helix of the (1 ! 4)-b-D-glucan, cellulose,
which has two hydrogen bonds (O20 H-O60 and O50 H-O30 )
between adjacent glucosyl units (Figure 1(b)). The absence of
cooperative interresidue associations in xylans allows the mol-
ecules to be conformationally more versatile. Indeed, in the
solid state, the unsubstituted (1 ! 4)-b-D-xylan chain preferen-
tially exists as a fully extended, threefold (i.e., three Xylp units
per helical turn), left-handed helix or a slowly twisted ribbon
217
(Figure 8(a)). Araf substitution of the backbone Xylp units
does not substantially change the basic threefold helical con-
formation (Figure 8(b) and 8(c)). However, substituted arabi-
noxylans cannot assume the twofold helical conformation in
solution because of significant steric interactions between
the Araf units and the xylan backbone. Nevertheless, (1 ! 4)-
b-D-xylans with low degrees of substitution interact reversibly
with the surfaces of cellulose microfibrils and may do so in
the twofold helical conformation. With increased Araf
substitution, the affinity for cellulose decreases, presumably
due to the shift from twofold to threefold helical conforma-
tion, which would be unfavorable for association with cellu-
lose, and because of the steric hindrance to interaction
imposed by the substituents themselves.
The length of the backbone chains of arabinoxylans is
 35 nm. In solution, the arabinoxylans behave as partially
stiff, wormlike, cylindrical molecules whose flexibility is largely
unaffected by Araf:Xylp ratios, which are generally in the range
0.390.82:1.
Solution Properties
The intrinsic viscosities of wheat arabinoxylans range from 0.8
to 5.5 dl g1, depending on the degree of substitution. These
values may be compared with 0.21 dl g1 for dextran, 0.19 for
beet arabinan, 0.120.25 for gum arabic, and 4.66.9 and
2.09.6 for barley and oat (1 ! 3, 1 ! 4)-b-D-glucans, respec-
tively. The behavior of the arabinoxylans in solution is influ-
enced not only by the overall asymmetry of the molecules and
their DP but also by the specific arrangement of Araf units
along the xylan backbone. Thus, stretches of unsubstituted
Xylp units may permit intermolecular alignment over these
sequences and lead to the formation of chain associations
that are stabilized by hydrogen bonds. As with (1 ! 3, 1 ! 4)-
b-D-glucans, the properties of arabinoxylan solutions are
strongly concentration-dependent. At low concentrations,
their viscosities are not shear-dependent but become so at
higher concentrations. This behavior is typical of polysaccha-
rides with extended, asymmetrical conformations. Removal of
Araf units, either by dilute acid or by a-L-arabinofuranosidase
action, leads to aggregation and precipitation of the residual
(1 ! 4)-b-D-xylan or of partially substituted arabinoxylan
molecules.
Gelation
At high concentrations, arabinoxylan solutions form thermo-
reversible gels that are stabilized by noncovalent interactions.
Gelation of arabinoxylan solutions can also be induced by
dimerization (cross-linking) of feruloyl units on neighboring
arabinoxylan chains by radical coupling (Figure 6). The dimer-
ization can be catalyzed by peroxidase/hydrogen peroxide,
manganese-dependent peroxidase, laccase, ammonium persul-
fate, ferric chloride, and other metal chlorides, halogens, and
linoleic acid/lipoxygenase, but not by iodate or bromate. Well-
developed, three-dimensional gel networks are obtained with
arabinoxylans containing high FA contents, high molecular
masses, and relatively unsubstituted xylan backbones. The lat-
ter facilitate the initial contact between feruloyl groups on
neighboring arabinoxylan chains. The rate of gelation depends
218 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides

Figure 8 Arabinoxylan conformations. Projections perpendicular (top) and parallel (bottom) of (a) (1 ! 4)-b-D-xylan backbone, (b) backbone with
single arabinofuranosyl side groups, and (c) backbone with two arabinofuranosyl side groups. Hydrogen bonds are shown by dotted lines. In each case,
the backbone is a left-handed, threefold helix. Reproduced from Atkins EDT (1992) Three dimensional structure, interactions and properties of xylans.
In: Visser J, Beldman G, Kusters-van Someren MA, and Voragen AGJ (eds.) Xylan and Xylanases, 576 pp. Amsterdam: Elsevier, with permission.

on temperature, pH, and concentration of oxidizing agent. The
gel-forming abilities of water-extractable arabinoxylans from
rye and barley are higher than for those from wheat and
triticale, presumably because of fine chemical differences of
the type listed earlier. Covalently cross-linked arabinoxylans
may hold up to 100 g water per 1 g polysaccharide. Gels
formed by treatment of maize bran with peroxide/peroxidase
are brittle, probably due to the high content of cellulose.
StructureFunction Relationships in Cereal Processing
and in Nutrition
Arabinoxylans, together with (1! 3, 1 ! 4)-b-D-glucans, con-
tribute to wort and beer viscosity, which may impede wort
separation and beer filtration. Arabinoxylans are also compo-
nents of some beer hazes. The water-soluble arabinoxylans in
barley, rye, and wheat contribute an antinutritive effect in
dietary formulations for poultry. In human diets, cereal arabi-
noxylans are components of the dietary fiber fraction. Water-
extractable fractions from rye, wheat, and spelt (Triticum spelta)
are mainly arabinoxylan, whereas those from barley and oats are
mainly (1! 3, 1 ! 4)-b-D-glucans. Ingestion of arabinoxylan-
enriched, wheat flour by-products reduces the glucose response
following a meal in normal subjects. Water-extractable cereal
fractions have a very low content of dehydrodiferulate and are
partially fermented by anaerobic bacteria in the colon to pro-
duce short-chain fatty acids. Water-unextractable cereal fractions
contain variable proportions of dehydrodiferulate; maize and
millet have the highest content and oat wheat and spelt the
lowest. Water-unextractable cereal fractions are poorly fermen-
ted in the colon, which may be related to the relatively high
diferulate cross-linking of arabinoxylan chains that prevents
swelling and attack by bacterial polysaccharide hydrolases or,
more generally, because of lower rates of pentose fermentation.
High concentrations of water-unextractable fractions in the diet
are effective in increasing stool weight, through its bulking
capacity. In recent pig feeding trial experiments, diets supple-
mented with wheat arabinoxylans were associated with
increased viscosity of small intestine digesta, changes in large
intestine microbiota composition that led to large increases in
 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
short-chain fatty acids, reduced damage to DNA from
colonocytes, and reduced serum triglyceride levels. These
changes can be linked with human health benefits that are
attributed to dietary fiber, including reduced glycemic index,
reduced risk of colorectal cancer and cardiovascular disease,
and reduced risk of type II diabetes.
Arabinoxylans have well-demonstrated impacts in bread
making. Added water-extractable arabinoxylans have positive
effects on the water absorption of doughs, especially after
oxidative gelation, and also enhance loaf volume. In both
cases, the amounts and molecular masses of the added arabi-
noxylan are important. Added water-extractable arabinoxylans
are reported to slow starch retrogradation and to produce less
firm breadcrumbs. The effect on breadcrumb texture is attrib-
uted to the increased moisture content of the samples. Water-
extractable arabinoxylans from wheat bran are good emulsion
stabilizers. They increase the strength and elasticity of gluten
starch films surrounding gas bubbles in doughs. This leads to a
higher retention rate of CO2 in the bubbles and produces a
positive effect on the fineness and homogeneity of crumb
texture. On the other hand, water-unextractable arabinoxylans
have detrimental effects on dough rheology and loaf volume.
Water-extractable rye arabinoxylans are inhibitors of ice crystal
growth and are thus candidate cryostabilizers. Arabinoxylan
lipid mixtures may be cast as films.
Analysis and Detection
The determination of arabinoxylans in cereal and cereal prod-
ucts relies on the phloroglucinolHCl or orcinolHCl colori-
metric reactions that show a high, but not absolute, specificity
for pentoses. In addition, NIR can be used to estimate arabi-
noxylan content in grain, and NIR peak intensity ratios at 1164
and 990 cm1 allow the degree of Araf substitution in arabi-
noxylans to be defined. Arabinoxylans in grain tissues can be
sensitively detected by UV-induced, bright blue fluorescence
(lmax, 415445 nm; excitation, 310 nm) due to esterified FA
residues. Lignins also fluoresce under UV irradiation but may
be distinguished from ferulate esters by examining walls at a
high pH, where the lignin continues to fluoresce blue, but
ferulate esters fluoresce green. Arabinoxylans may also be
located in tissue sections through immunogold labeling with
polyclonal antibodies raised against 4-O-methyl glucuronoxy-
lan or arabinoxylan or with gold-labeled xylanases. Polyclonal
antibodies raised specifically against highly or sparsely
substituted glucuronoarabinoxylans can also differentiate
these types of arabinoxylan in cell walls.
Genetics
As high-density genetic maps for barley and wheat have
become available, QTLs for arabinoxylan (or pentosan) con-
tent in grain have been mapped. A QTL accounting for  35%
of variation in both Araf: Xylp ratio and extract viscosity has
been found on the long arm of chromosome 1B of bread
wheat. The observation that there is no apparent change in
arabinoxylan concentrations in the wheat 1BL/1RS transloca-
tion lines suggests that the genes involved are located outside
the segment of that chromosome that was introgressed from
rye. A total of five QTLs controlling acid detergent fiber (ADF)
219
Table 5 Genes that have been implicated in heteroxylan biosynthesis
in higher plants, including proposed roles in xylan backbone synthesis
and the addition of the numerous substituents of the backbone chain
Gene family Proposed role
GT8 (IRX8 and GAUT) GluA transferases
GT43 (IRX9 and IRX14) (1,4)-b-Xylan synthase
GT47 (IRX10 and IRX7) (1,4)-b-Xylan synthase
GT61 Ara transferases or Xyl transferases
BAHD/PMT acyltransferases Feruloyl transferases
DUF579 (IRX15) O-Methyl transferase (GluA)
AXY O-Acetylation
have been identified in barley, and although ADF consists pre-
dominantly of cellulose and lignin from the pericarpseed coat,
some of the effects might be related to arabinoxylan and/or
(1! 3, 1 ! 4)-b-D-glucan content. Three QTLs, with relatively
large effects, map close to one another on chromosome 2H,
whereas the other two QTLs are on chromosomes 4H and 1H.
Overall, however, interpretation of the QTL data in terms
of identifying genes involved in the biosynthesis of arabinox-
ylans has usually been complicated by the phenotyping
assays. For example, QTLs for water-extractable arabinoxylans
might be attributable either to xylan synthase activity or to
arabinofuranosidase/glucuronidase activity, because solubil-
ity will be affected by the degree of substitution of the xylan
backbone. Phenotyping on the basis of Araf or Xylp levels or
Araf:Xylp ratios can also be misleading, because both mono-
saccharides will be found in polysaccharides other than ara-
binoxylans, such as xyloglucans, pectin, and AGPs.
Based largely on the analyses of Arabidopsis mutants, a
number of genes have been implicated in heteroxylan biosyn-
thesis (Table 5). However, the roles of these genes in arabinox-
ylan synthesis in cereal grains have not always been defined
and the identification of genes that mediate the biosynthesis of
the (1 ! 4)-b-D-xylan backbone of cereal arabinoxylans still
requires confirmation.
Glucomannans
Glucomannans are linear copolymers of b-D-glucopyranose
( 30%) and its 2-epimer, b-D-mannopyranose ( 70%),
joined by (1 ! 4)-linkages (Figure 9(a)) to form linear chains
with DPs ranging from < 100 to several thousand. In some
examples, the backbone has (1 ! 6)-linked b-D-
galactopyranosyl substituents and is generally esterified with
acetyl groups. The shape of the glucomannan polymer is sim-
ilar to cellulose (Figure 1(a)), and it has therefore been sug-
gested that the chains may associate strongly with surfaces of
cellulose microfibrils. The glucomannans are extractable from
walls with alkaline borate solutions that act by complexing
with the mannopyranosyl units.
Glucomannans are minor constituents of walls of most
aleurone and starchy endosperm cells of cereals (Table 1),
but in certain Oryza indica rice cultivars, values of up to 17%
glucomannan have been reported in the endosperm walls. The
CslA genes have been shown to mediate glucomannan biosyn-
thesis in some plants.
220 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides

Gal
6
--Man 4 Man 4 Glc 4 Man 4 Man 4 Man 4 Man 4 Man 4 Man 4 Glc 4 Glc 4 Man--
6
Gal
Man = -D-mannopyranosyl unit
Glc = -D-glucopyranosyl unit
Gal = a-D-galactopyranosyl unit
4 = (14)-linkage
(a) 6 = (16)-linkage

Fuc Fuc R R R
2 2 4 4 4
Gal Gal --4 Gal A 2 Rha 4 Gal A 2 Rha 4 Gal A 2 Rha 4--
2 2
Gal A = -D-galaclopyranosyl unit
Xyl Xyl Xyl Xyl
Rha = a-L-rhamnopyranosyl unit
6 6 6 6
4 = (14)-linkage
--Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc 4 Glc--
2 = (12)-linkage
6 6
R = (14)-linked -D-galactopyranosyl chain
Xyl Xyl
or
Glc = -D-glucopyranosyl unit (13,16)-linked -D-galactopyranosyl chain
Xyl = a-D-xylopyranosyl unit or
Gal = -D-galactopyranosyl unit (15)-linked a-L-arabinofuranosyl chain with
Fuc = a-L-fucopyranosyl unit (c) (12) and (13)-side-branching
4 = (14)-linkage
6 = (16)-linkage
(b) 2 = (12)-linkage

Figure 9 Structural features of three noncellulosic polysaccharides found as minor constituents in cell walls of cereal grains: (a) glucomannans,
(b) xyloglucans, and (c) pectic polysaccharides.

Xyloglucans There is also variable methyl esterification and hydroxyl acety-

Xylo-(1 ! 4)-b-D-glucans (xyloglucans) are a family of side
chain-branched heteroglycans (Figure 9(b)) consisting of a
cellulose-like (1 ! 4)-b-D-glucan backbone side-branched at
regular intervals by a-D-xylopyranosyl units at C(O)6 of the
glucosyl residues. Some side-branch xylosyl units carry b-D-
galactopyranosyl and a-L-fucopyranosyl substituents, and the
unsubstituted glucosyl residues of the backbone chain can be
acetylated. Compared with xyloglucans from dicotyledonous
plants, those from the cereals are less substituted with xylose,
contain little galactose, and have stretches of unsubstituted
backbone. Xyloglucans have DPs of 600700 and readily asso-
ciate with surfaces of cellulose microfibrils. After extraction
with dilute alkali, they are water-soluble. Xyloglucans are gen-
erally considered to be of minor significance in walls of cereal
grain tissues, although procedures used in earlier analyses may
have underestimated their relative abundance in wall
preparations.
Pectic Polysaccharides
The pectic polysaccharides are a diverse group of side chain-
branched heteroglycans. The backbone is a copolymer of alter-
nating (1 ! 4)-linked a-galacturonosyl and (1! 2)-linked
a-L-rhamnopyranosyl units (Figure 9(c)). The rhamnogalactur-
onans are variously side-branched on the rhamnosyl units by
arabinan, arabinogalactan, and more complex branched oligo-
saccharides. In addition, variable amounts of (1 ! 4)-a-linked
homogalacturonan may be present in the same backbone chain.
lation of the galacturonosyl units. The homogalacturonan
domain complexes with Ca2, which promotes gelation by
forming Ca2 bridges between unesterified galacturonosyl
units on adjacent chains. Pectic polysaccharides are minor con-
stituents of the walls of vegetative tissues and grains of cereals,
although measurable amounts have been reported in the walls
of rice endosperm cells (Table 1).
Callose
Callose is a (1 ! 3)-b-D-glucan. Like cellulose, it is a linear
homopolymer and is insoluble in water. However, callose is
soluble in dilute NaOH. Callose can be recognized in tissue
sections by an intense yellow, UV-induced fluorescence (lmax,
495 nm; excitation, 380 nm) when associated with the aniline
blue fluorochrome, a sulfonated 4,40 -dianilinobenzophenone,
or by its binding to a specific monoclonal antibody. In mature
barley endosperm, callose occurs as small bead-like deposits
on the inner surfaces of walls throughout the starchy endo-
sperm but especially at the aleurone/subaleurone interface.
The deposition of callose is a well-known consequence of stress
and wounding. Thus, the endosperm deposits may arise as a
consequence of plasmolysis during the drying of the grain. The
callose content of barley endosperm wall preparations is  1%.
Callose is found in the developing walls of rice and barley
endosperm during the early stages of cellularization, but is
not a component of mature walls. It is also found in the
nucellar projection and in the vascular tissue of the crease of
developing barley but at maturity remains only in the vascular
 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides 221

tissue. In the nucellar projection and the vascular tissue, callose specific enzymatic procedure, and their size distribution deter-
deposition might be related to the control of assimilate trans- mined by HPLC or MALDI-TOF-MS.
port into the developing endosperm.

Fructans
Fructans from cereals consist of short chains of up to 45 b-D-
fructofuranosyl units in (2 ! 6)-b-linkage to the fructosyl res-
idue of the disaccharide sucrose. Some branching through
(2 ! 1)-b-linkages may also occur. Cereal fructans are soluble
in water and in boiling 80% (v/v) ethanol. During the early
development of wheat grain, fructans can account for as much
as 20% dry weight. In mature grains, the contents are low and
variable: wheat, 1.32.5%; rye, 4.66.6%; barley, 0.8%; and
oats, 0.1%. In the grain, fructans are found in both the endo-
sperm and the pericarp/embryo, with higher concentrations in
the latter fraction.
Fructans are important carbohydrate reserves in the vegeta-
tive tissues of temperate cereals, particularly in young seedlings
where they can account for 70% of the dry weight. Cereal
fructans are not digested in the monogastric stomach or small
intestine but are readily fermented by microorganisms in the
lower gut. Thus, they may be considered to be components of
cereal dietary fiber. Fructans may be quantified by NIR or by a
Arabinogalactan Proteins
AGPs are widely distributed in plants. They consist of a protein
backbone that is rich in hydroxyproline and to which arabino-
galactan polysaccharides are covalently attached, although
there are many structural subclasses, some of which are
membrane-anchored. In wheat grains, the major AGP is an
arabinogalactan peptide, with a peptide backbone that is
much smaller than most AGPs. The arabinogalactan peptide
from wheat flour constitutes 0.270.38% on a dry-weight basis
and has an average molecular mass of 22 000, although higher-
molecular-mass AGPs have been detected in other cereal
grains, including barley. The polysaccharide portion of the
wheat arabinogalactan peptide constitutes 92% of the
molecule and consists of branch-on-branch b-D-galactosyl
units (Galp) linked by (1 ! 3)- and (1 ! 6)-linkages
(Figure 10). The branched b-D-galactan backbone is
substituted by single a-L-arabinofuranosyl units. The arabino-
galactan has an Araf:Galp ratio of 0.630.72:1. In wheat endo-
sperm, the arabinogalactan component of the molecule is
covalently linked to a peptide of molecular mass 7800 at

+
H3N Tyr Ala Glu Val Hyp Ser Hyp Ala Ala Gly Ala Hyp Thr Ala Asp COO

Figure 10 Structure of wheat arabinogalactan peptide. () (1 ! 3)-Linked-b-D-galactopyranosyl unit, ( ) (1 ! 6)-linked b-D-galactopyranosyl unit,
and ( ) terminal a-l-arabinofuranosyl unit. Structure based on data of Fincher GB, Sawyer WH, and Stone BA (1974) Chemical and physical
properties of an arabinogalactan-peptide from wheat endosperm. Biochemical Journal 139: 535545; Van den Bulck K, Looseveld A-MA, Courtin CM,
Proost P, Van Damme J, Robben J, Mort A, and Delcour JA (2002) Amino acid sequence of wheat flour arabinogalactan-peptide, identical to part
of grain softness protein GSP-1 leads to improved structural model. Cereal Chemistry 79: 329331.
222 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
each of the three hydroxyproline residues in the peptide
sequence. The same peptide sequence is found at the NH2-
terminus of a protein associated with the surface of wheat
starch granules. Among eight Canadian wheat cultivars, the
peptide constituted between 6.5% and 14.3% of the AG-
peptide molecule.
The wheat AG peptide is very soluble in water and is not
precipitated at saturating ammonium sulfate concentrations.
Its solutions are of low intrinsic viscosity (0.0450.062 dl g1).
Neither its subcellular location in the endosperm nor its phys-
iological function is known. The wheat AG peptide, unlike its
larger AGP counterparts, does not bind to the phenylazo dye,
b-glycosyl Yariv reagents. The effects of wheat AG peptides on
bread-making properties are of equivocal significance.
Concluding Remarks
There is a great deal of information available on the chemistry
and physicochemistry of nonstarchy polysaccharides in cereal
grains. This is directly attributable to the central importance of
these polysaccharides in large-scale food processing activities
that include brewing, baking, and stock feed manufacture.
Moreover, the nonstarchy polysaccharides of cereals have
attracted renewed interest in recent years because of their
potentially beneficial effects in human nutrition. Despite this
interest, there remain major gaps in our knowledge of the
genes and enzymes that control nonstarchy polysaccharide
biosynthesis in the cereal grain, particularly the genes that
control arabinoxylan biosynthesis. In the case of cereal
(1 ! 3, 1 ! 4)-b-D-glucans, where genes involved in the bio-
synthesis of the polysaccharide have been identified, attention
is now turning to genes that regulate levels of grain (1 ! 3,
1 ! 4)-b-D-glucan and eventually to an understanding of the
factors that control fine chemical structure of the polysaccha-
rides and hence their solubility and general solution behavior
during cereal processing. This information can be confidently
expected to provide additional molecular markers through
which breeders will track key genes in their breeding programs,
to allow breeders to improve quality characteristics related to
nonstarchy polysaccharides in commercially important pro-
cesses, and to further enhance the role of cell wall polysaccha-
rides as components of dietary fiber that carry human health
benefits.
Exercises for Revision
What differences would you expect in the solubility of
wheat flour arabinoxylans with xylosearabinose ratios of
1.2 and 1.5, respectively, and how would you rationalize
those differences in terms of chemical fine structure?
What differences would you expect in the solubility of
barley (1 ! 3, 1 ! 4)-b-D-glucans with DP3:DP4 ratios of
2.5 and 3.0, respectively, and how would you rationalize
those differences in terms of chemical fine structure?
Cereal grains produce noncellulosic cell wall polysaccha-
rides, such as arabinoxylans and (1 ! 3, 1 ! 4)-b-D-glucans,
that will not form microfibrils of the type produced by
cellulose. What are the two different chemical strategies
used to prevent microfibril formation in arabinoxylans
and (1 ! 3, 1 ! 4)-b-D-glucans?
Summarize current knowledge on the genes that mediate
arabinoxylan and (1 ! 3, 1 ! 4)-b-D-glucan biosynthesis in
cereal grains.
Summarize the functions of arabinogalactan peptides in
wheat grains.
Show the detailed metabolic pathways leading to the struc-
tures shown in Figure 7.
Exercises for Readers to Explore the Topic Further
Although there have been significant advances in our under-
standing of the metabolism of cereal grain nonstarchy poly-
saccharides, there remain a number of unanswered questions
that will undoubtedly attract research attention in the imme-
diate future. Examples of these key questions include the
following:
What is the precise molecular mechanism through which
(1 ! 3, 1 ! 4)-b-D-glucans are synthesized and how are the
fine structures and hence physicochemical properties
altered in response to changing functional requirements?
What genes are involved in the biosynthesis of arabinoxy-
lans in cereal grains?
Can we identify regulatory genes that control and amounts
and fine structures of (1 ! 3, 1 ! 4)-b-D-glucans and arabi-
noxylans in cereal grains?
Can we define the cellular aspects of (1 ! 3, 1 ! 4)-b-D-
glucans and arabinoxylan synthesis, including sequential
assembly events and the organelles involved?
Is there evidence for the two phase assembly of cell wall
polysaccharides, whereby oligosaccharides are synthesized
first and subsequently assembled into polysaccharides?
See also: Carbohydrates: Health Effects of b-Glucans Found in
Cereals; Food Grains and the Consumer: Grains and Health; The
Basics: Grain: Morphology of Internal Structure; The Cereal Grains:
Barley: An Overview of a Versatile Cereal Grain with Many Food and
Feed Uses; Wheat: An Overview of the Grain That Provides Our Daily
Bread.
Further Reading
Atkins EDT (1992) Three-dimensional structure, interactions, and properties of xylans.
In: Visser J, Beldman G, Kusters-van Someren MA, and Voragen AGJ (eds.) Xylan
and Xylanases, p. 576. Amsterdam: Elsevier.
Bamforth CW (1994) b-Glucans and b-glucanases in malting and brewing: Practical
aspects. Brewers Digest 69: 1216.
Belobrajdic DP, Bird AR, Conlon MA, et al. (2011) An arabinoxylan-rich fraction from
wheat enhances caecal fermentation and protects colonocyte DNA against diet-
induced damage in pigs. British Journal of Nutrition 107: 19.
Burton RA, Collins HM, Kibble NAJ, et al. (2011) Over-expression of specific HvCslF
cellulose synthase-like genes in transgenic barley increases the levels of cell wall
(1,3;1,4)-b-D-glucans and alters their fine structure. Plant Biotechnology Journal
9: 117135.
Burton RA, Gidley MJ, and Fincher GB (2010) Heterogeneity in the chemistry, structure
and function of plant cell walls. Nature Chemical Biology 6: 724732.
Burton RA, Wilson SM, Hrmova M, et al. (2006) Cellulose synthase-like CslF genes
mediate the synthesis of cell wall (1,3;1,4)-b-D-glucans. Science 311: 19401942.
 CARBOHYDRATES | Cereals: Chemistry and Physicochemistry of Non-starchy Polysaccharides
Carpita N and Gibeaut D (1993) Structural models of primary cell walls in flowering
plants: Consistency of molecular structure with the physical properties of the walls
during growth. Plant Journal 3: 130.
Collins HM, Burton RA, Topping DL, Liao M-L, Bacic A, and Fincher GB (2010)
Variability in the fine structures of non-cellulosic cell wall polysaccharides from
cereal grains: Potential importance in human health and nutrition. Cereal Chemistry
87: 272282.
Cui SW (2001a) Cereal non-starch polysaccharides I: (1 ! 3) (1 ! 4)-b-D-glucans.
In: Cui SW (ed.) Polysaccharide Gums from Agricultural Products, pp. 104165.
Lancaster: Technomic Publishing.
Cui SW (2001b) Cereal non-starch polysaccharides II: Pentosans/Arabinoxylans.
In: Cui SW (ed.) Polysaccharide Gums from Agricultural Products, pp. 167227.
Lancaster: Technomic Publishing.
Evers T and Millar S (2002) Cereal grain composition and development: Some
implications for quality. Journal of Cereal Science 36: 261284.
Fincher GB, Sawyer WH, and Stone BA (1974) Chemical and physical properties of an
arabinogalactan-peptide from wheat endosperm. Biochemical Journal 139: 535545.
Fincher GB and Stone BA (1986) Cell walls and their components in cereal grain
technology. Advances in Cereal Science and Technology 8: 207295.
Grabber JH, Hatfield RD, Ralph J, Zon J, and Amrhein N (1995) Ferulate cross-linking
in cell walls isolated from maize cell suspension cultures. Phytochemistry
40: 10771082.
223
Harris PJ and Fincher GB (2009) Distribution, fine structure and function of (1,3;1,4)-b-
glucans in the grasses and other taxa. In: Stone BA, Bacic T, and Fincher GB (eds.)
Chemistry, Biochemistry, and Biology of (1,3)-b-Glucans and Related
Polysaccharides, pp. 621654. Philadelphia, PA: Elsevier.
Iiyama K, Lam TB-T, and Stone BA (1994) Covalent cross-links in the cell wall. Plant
Physiology 104: 315320.
Izydorczyk MS and Biliaderis CG (1995) Cereal arabinoxylans Advances in structure
and physico-chemical properties. Carbohydrate Polymers 28: 3348.
Kroon-Batenburg LMJ and Kroon JJ (1995) The crystal and molecular structures of
cellulose. Carbohydrates in Europe 12: 1519.
Skendi A, Biliaderis CG, Lazaridou A, and Izydorczyk MS (2003) Structure and
rheological properties of water soluble b-glucans from oat cultivars Avena sativa
and Avena bysantina. Journal of Cereal Science 38: 117.
Stone BA and Clarke AE (1992) The Chemistry and Biology of (1 ! 3)-b-Glucans.
Victoria: Latrobe University Press.
Van den Bulck K, Looseveld A-MA, Courtin CM, et al. (2002) Amino acid sequence of
wheat flour arabinogalactan-peptide, identical to part of grain softness protein GSP-
1 leads to improved structural model. Cereal Chemistry 79: 329331.
Vinkx CJA and Delcour JA (1996) Rye (Secale cereale L.) arabinoxylans A critical
review. Journal of Cereal Science 24: 114.
Visser J, Beldman G, Kusters-van Someren MA, and Voragen AGJ (eds.) (1992) Xylan
and Xylanases, p. 576. Amsterdam: Elsevier.
 Grains other than Cereals: Non-starch Polysaccharides
L Ramsden, University of Hong Kong, Hong Kong, China
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Non-starch polysaccharides (NSP) have a diverse composi-
tion and are widely distributed in plants, are all
indigestible, but can strongly influence texture.
Different types of NSP are produced in monocots and
dicots.
NSP may be accumulated as a reserve polysaccharide inde-
pendently or as part of the cell wall.
NSP may influence the absorption of carbohydrates, lipids,
and proteins.
The legume family is an important source of non-cereal
grains.
Learning Objective
To appreciate the diversity, abundance, and potential value
of NSP resources in non-cereal grains.
Introduction
All plant tissues contain a large variety of polysaccharides,
many of which serve structural or reserve functions in the
plant. However, the most important polysaccharide obtained
from grain crops is starch and the focus in starch producing
grain crops is naturally on the yield of starch. Any other poly-
saccharides present are considered to be of secondary impor-
tance as non-starch polysaccharides (NSPs), a term that covers
a great variety of biological functions and chemical structures.
There is a growing awareness that these polysaccharides can
have considerable value both in human nutrition and for the
food industry. In particular, non-cereal grains show an inter-
esting diversity of polysaccharides that are less familiar than
the better studied NSPs of the cereal grain. Non-cereal grains
include a great variety of potential species, many of which have
not been analyzed for NSP content. Consequently here, it is
only possible to present a limited selection together with some
details of the structure and their value in products derived from
grains.
Polysaccharides in a grain can serve two major biological
functions. They can act as the energy reserve in the endosperm
or cotyledons or they may be involved in forming the structure
of the grain tissues. Other functions in which polysaccharides
can play essential roles are in the regulation of water balance
during dormancy and imbibition and in the protection of the
seed against predation and attack by pathogens. In cereals,
only starch has a major reserve function, but in dicot grains,
other polysaccharides are found to act as reserves, often
together with starch.
224 Encyclopedia of Food Grains, Second Edition
Plant Cell Walls
In common with all plant tissues, the tissues forming a grain
are composed of cells, each of which is defined by a cell wall.
The cell is the basic structural unit of plants and cellulose is the
basic structural component of the plant cell wall. All plant cell
walls contain not only cellulose but a number of other poly-
saccharides that are important in maintaining the structural
integrity of the wall and controlling wall permeability. Other
components may be involved in interactions with neighboring
cells and the environment.
The cell walls of plants display a composition that is charac-
teristic of the taxonomic group in which the plant is found. In
the angiosperms, there are consistent differences in the pattern
of wall polysaccharides present between dicots and monocots.
Thus, cereal plant cells possess walls typical of monocot grasses,
and this also applies to the endosperm cells in the cereal grain.
The cell wall is formed from two types of wall, the primary and
the secondary. Primary walls are those first laid down after a cell
is formed and are thinner and more flexible than secondary
walls that are found in mature cell types. Both primary and
secondary walls contain cellulose, but the cellulose fibrils of
the primary wall are thinner and more loosely arranged; in the
secondary wall, the cellulose fibrils are closely packed into
organized layers that are linked to each other by cross-links.
Endosperm cells are typically larger and have thinner walls
than other tissues in the plant so the proportion of cellulose
present can be lower than that in other tissues.
Seed-bearing plants deposit energy-containing reserves to
support growth of the embryo within the seed. While the major
reserve polysaccharide is indisputably starch, there are a num-
ber of other polysaccharides that are used particularly within
the legume plant family (Fabaceae). Starch reserves are stored
in the cytoplasm inside amyloplasts, but the non-starch reserve
polysaccharides in seeds are deposited in highly thickened
walls of endosperm cells without a structural function.
Distribution of Grain NSPs
There is a clear distinction between monocot- and dicot-grain
NSPs, which reflects the differences between the monocot and
dicot cell walls. The actual levels present in any grain sample
can also vary dependent on the growth conditions and partic-
ular genotype. The distribution of NSPs in dicot grains is more
difficult to assess due to the variability in designation of what
constitutes a dicot grain. In dicots, the major monocot NSPs,
the mixed linkage b-glucans, are absent (Figure 1), and arabi-
noxylans, though they may be present in dicot cell walls,
should generally be expected to be largely replaced by xyloglu-
can that is the main hemicellulose in dicots. Xyloglucans can
be found as a reserve polysaccharide in a number of seeds.
Galactomannans are found as seed reserve polysaccharides in
http://dx.doi.org/10.1016/B978-0-12-394437-5.00091-7
 CARBOHYDRATES | Grains other than Cereals: Non-starch Polysaccharides 225

14 glucose 13 glucose

Figure 1 The sequence of glucose units in a mixed linkage (13)(14)-b-glucan.

several dicot families; though their commercial importance as Table 1 Dietary fiber in cereal and legume grains
food additives is derived from their presence in large tree seeds,
they are also present in many smaller grains. Total fiber

Wheat 14.6
Uses of Grain NSPs Barley 22.6
Rice 2.4
Sorghum 10.1
The presence of high levels of NSPs in grain products will
Millet 8.5
reduce the nutritional value of the grain, a factor which is of
Maize 13.4
significance in the animal feed industry. An important feature Oats 9.6
of NSPs is their ability to bind water that allows them to act as Rye 14.6
hydrocolloids and disproportionately influence the rheology Triticale 18.1
of aqueous systems. In baking, the NSPs of wheat can enhance Chickpea 13.5
water retention and texture of bread and cakes. Purified NSPs Black gram 7.1
can be used as raw materials for the food additive industry that Mung bean 12.8
is a major user of polysaccharide texture modifiers, stabilizers, Pigeon pea 8.1
and gelling agents. Currently, traditional grains are not major Soybean 4.96
Lupin 4.0
sources of industrial polysaccharide hydrocolloids, but high
Lima bean 6.3
demand on traditional supplies makes this a competitive area
Kidney bean 6.2
for development of new sources. However, one of the major Faba bean 8.0
benefits of NSPs arises after they have been consumed as part Sunflower 4.2
of the diet, when they contribute to the dietary fiber.
Figures given are the highest levels reported. Actual values may vary considerably
depending on the variety grown and the cultivation conditions.
Dietary Fiber Data from Shelton DR and Lee WJ (2000). In: Kulp K and Ponte JG (eds.) Handbook of
Cereal Science and Technology, 2nd edn., pp. 385415. New York: Marcel Dekker;
The NSPs together form the major part of the dietary fiber of Sathe SK (1996). In: Nwokolo E and Smartt J (eds.) Food and Feed from Legumes and
grain crops. Dietary fiber is the fraction of a food consumed that Oilseeds, pp.1232. London: Chapman & Hall.
is not degraded in the small intestine, and as human digestive
enzymes can only cleave a14 glucan bonds, all polysaccharides less well suited to the modern high-fat, energy-rich, low-volume
other than starch are part of the dietary fiber. Dietary fiber can be diets of industrialized countries. This deficiency in the modern
both soluble and insoluble dependent on the molecular struc- diet can be overcome through the consumption of grain prod-
ture of the polysaccharides. There are also a smaller number of ucts high in dietary fiber. NSP can lower the availability and
other polymers present in plant tissues, such as lignin, which glycemic index of starch by complex formation and restricting
also form part of the dietary fiber. However, the simplest avail- access for digestive enzymes. Polysaccharides can also bind to
able measure of the quantity of NSP in grains is the level of dietary lipids and reduce their uptake into the body, a factor that
dietary fiber present (Table 1), and often, this is the only figure has been observed in the ability of high-fiber diets to lower
available for many grains that have not been analyzed in ade- blood lipid levels with potentially beneficial consequences for
quate detail for polysaccharide composition. the incidence of cardiovascular disease.
Although unable to contribute to human nutrition in terms
of provision of energy, the dietary fiber is well recognized to
form an important component of our diet for the correct func-
tioning of the digestive system. Polysaccharides through their Insoluble Cell Wall Polysaccharides
high water-binding capacity play an important part in providing
bulk to the gut contents to allow easy passage through the The main structural component of any cell wall is cellulose, a
intestine. The human digestive system developed to cope with b14 linked polymer of glucose (Figure 2). It is the worlds most
a diet rich in high fiber plant material with a large volume and is abundant polymer, followed by its sister glucan starch, with the
226 CARBOHYDRATES | Grains other than Cereals: Non-starch Polysaccharides
14 glucose
Figure 2 The sequence of sugar units in the b14 glucan cellulose.
key difference that it cannot be degraded by human digestive
enzymes and cannot therefore contribute directly to our nutri-
tion. Cellulose chains are long flat linear ribbons of glucose
units, the number of which can exceed 10 000 and with molec-
ular weights of over 1 000 000. Because the b14 linkage between
the glucose units holds the chain in a flat conformation, it is
possible for cellulose chains to align next to each other and form
numerous hydrogen bonds between the sugar hydroxyl groups.
The chains can stack together to form larger microfibrils, which
makes cellulose highly insoluble and tough, an ideal building
material for plants. The quantity of cellulose that is found in
grains can vary from species to species and is largely a conse-
quence of the thickness of the husk and seed coat that tends to
have stronger thicker cell walls that contain more cellulose. The
cells of seed endosperm contain only thin cell walls, and in a
well-filled grain, the proportion of cellulose to starch or other
reserve polysaccharides should be low. Other insoluble polysac-
charides such as glucomannans can occur in the walls of some
plants, but quantities are typically low, and although levels of up
to 2% in some grain walls have been indicated, these would not
have a major impact on grain properties.
Soluble Cell Wall Polysaccharides
All plant cell walls contain more soluble polysaccharides
known as pectins and hemicelluloses with a variety of different
structures that serve various functions. These polymers contain
a number of different sugar units and are classified according
to their composition and solubility. The pattern of soluble
polysaccharides present is characteristic of groups and species
of plants.
The pectins are a major class of highly soluble wall-related
polysaccharides that are accumulated in fruits but not to a great
extent in most seed grains. Pectins are charged, acidic poly-
saccharides with a variety of structures based on rhamnogalac-
turonans, polymers of rhamnose and galacturonic acid
together with a number of other sugar substituents.
Purified hemicelluloses can show varying degrees of water
solubility dependent on their size and structure, but all hemi-
celluloses are strongly bound to the intact cell wall either by
hydrogen bonding or by cross-linking to cellulose. The major
function in the plant of hemicelluloses is to increase the rigid-
ity and impermeability of the cell wall.
Dicot Arabinoxylans
Arabinoxylans can be found in many dicots but usually only at
low levels. An exception is in linseeds, the seeds of the flax
plant (Linum usitatissimum). Linseed is a traditional crop val-
ued for the oil that can be expressed from the seeds. The seeds
readily hydrate to form an arabinoxylan mucilage that can
easily be extracted by incubation in cold water.
High levels of arabinoxylans are known to occur in seeds
of various species of genus Plantago. The arabinoxylan is
present in the seed coat and can be extracted with boiling
water and yields are increased under mild alkaline conditions.
The structure of the psyllium arabinoxylan is based on a xylan
backbone substituted with arabinose and some uronic acids
(Figure 3). Solubility in water is not high and the polysaccha-
ride swells to give a weak gel. The gel shows a broad melting
range around 80 and is susceptible to syneresis on freezing/
thawing but shows good stability over a range of ionic concen-
trations. Psyllium arabinoxylan is a soluble dietary fiber that is
effective in lowering plasma cholesterol levels and that has
traditionally been a component in laxatives where its high
swelling power and mucilaginous gel ease the passage of gut
contents.
Another abundant arabinoxylan is obtained from seeds of
the quince tree (Cydonia oblonga). Seeds can be directly
extracted with hot or cold water, and a mucilage is readily
released that is a mixture of cellulose microfibrils dispersed in
a matrix composed principally of xylose and arabinose. The
polysaccharide is easily soluble in cold water and forms a
highly viscous mucilage but is not gel-forming. Dispersions
show good stability with respect to pH, salt concentration,
and temperature. Quince seed arabinoxylan is mainly used
for cosmetics applications in Middle Eastern countries. A num-
ber of food uses have been suggested such as for use as a
stabilizer in ice cream, but these have not been commercialized
due to the high cost and poor availability and variable quality
of the gum.
Dicot Xyloglucans
The major hemicellulose of the dicot cell wall is a xyloglucan
formed from a b14 linked chain of glucose units substituted
with a16 xylose units. Some of the xylose units may be joined
to an a12 linked galactose unit (Figure 4). Most legume
grains contain a major proportion of xyloglucan in the cell
wall as a structural hemicellulose. Less often high levels of
xyloglucans are found in the seeds of some plants where they
appear to have a reserve function such as in nasturtium (Tro-
paeolum majus) and the tamarind tree (Tamarindus indica).
Tamarind seeds can be ground to flour that is hot water-
soluble to form a mucilaginous gel. The main polysaccharide is
a xyloglucan with a b14 glucan backbone carrying 16 linked
xylose and arabinose and galactose substituents in a ratio of
galactose/xylose/glucose of approximately 1:2:3. Tamarind xylo-
glucan can form a gel under acid conditions, which remains stable
at alkaline pH. Gels can also form in the presence of ethanol
where cross-links arise due to the low solubility of some chain
regions allowing aggregates to form. Tamarind gels are used in
India for a range of traditional confectionery products. Xyloglu-
cans are used as food additives in Japan for a wide range of
products.
 CARBOHYDRATES | Grains other than Cereals: Non-starch Polysaccharides
14 Xylose
-Arabinose
Figure 3 The arrangement of sugar units in arabinoxylan. a-Arabinose and a-glucuronic acid linked 12 or 13 to a backbone chain of b14 xylose units.
14 glucose
Xylose
Figure 4 The arrangement of sugar units in xyloglucan. a-Xylose linked 16 to a backbone chain of b14 glucose units, with some b-galactose
linked 12 to the xylose units.
Reserve Polysaccharides
Seed-bearing plants deposit energy-containing reserves to sup-
port growth of the embryo within the seed. While the major
seed reserve polysaccharide is without doubt starch, there are a
number of other polysaccharides that are used as reserves,
particularly within the legume family. Unlike starch, which is
stored in amyloplasts in the cytoplasm, the major non-starch
reserve polysaccharides in seeds are deposited in the cell wall
but do not serve a structural function.
Apart from starch, there is only one other major polysaccha-
ride used by plants as a reserve outside the cell wall. These are
polymers of fructose, the fructans, which can have chain lengths
of up to 250 sugar units but are generally shorter and highly
water-soluble. Many grasses are known to use fructans for stor-
age and for frost resistance. Potentially, fructans may occur in
the grains but there is little evidence for significant quantities
and most sources of fructans are from tubers or stem tissue.
Galactomannans
Many seeds of the legume family contain high levels of reserve
galactomannans. The seed galactomannans all possess the
same basic structure of an a14 linked mannan backbone
chain with varying degrees of substitution with a16 linked
227
-Glucuronic acid
Galactose
galactose residues (Figure 5). The ratio of galactose to man-
nose is roughly constant for a given species. The distribution of
galactose along the mannan chain is not uniform but tends to
be clustered in blocks of high substitution (rough regions) that
are separated by intervening stretches with few galactose resi-
dues (smooth regions).
Purified galactomannans for industrial usage are obtained
from four main plant sources in the legume subfamily Ca-
esalpinioideae that are well characterized. There are guar gum
from the seeds of Cyamopsis tetragonoloba, locust bean gum
from seeds of Ceratonia siliqua (carob tree), tara gum from
seeds of Caesalpinia spinosa, and cassia gum from seeds of
Cassia obtusifolia. Extraction of galactomannans involves de-
hulling of seeds and crushing to remove the embryo, followed
by milling of the endosperm to produce a crude flour. The
flour can be purified by dissolving in hot water, followed by
filtration and precipitation with isopropanol to remove impu-
rities. The properties of galactomannans from different sources
vary, depending on the structure of the galactomannan that is
characteristic of the source species.
The use of locust bean galactomannan (LBG) in the Medi-
terranean and the Middle East has been part of traditional food
preparation for hundreds of years. LBG has a galactose to
mannose ratio of 1:4 and a molecular weight of around
300 000. The cold water solubility of LBG is low and disper-
sions need to be heated to 85  C to achieve good dissolution,
228 CARBOHYDRATES | Grains other than Cereals: Non-starch Polysaccharides
14 mannose
16 galactose
Figure 5 The arrangement of sugars in galactomannan. a-Galactose
linked 16 to a backbone chain of b14 mannose units.
concentrations of up to 5% w/v being possible. Low solubility
is due to the tendency for the linear mannan chains to strongly
hydrogen bond to each other in unsubstituted regions of the
chain, limiting opportunities for interaction with water
molecules.
Though LBG does not itself form gels, it can be used
together with other hydrocolloids to give rise to gels, such
synergistic gel formation being observed with other non-
gelling polysaccharides such as xanthan gum. This property is
attributed to the ability of the nonsubstituted regions of the
linear mannan backbone being able to hydrogen bond to
helical regions of the other hydrocolloids and generate cross-
linking. LBG is widely used as a thickener and stabilizer in
many foods such as ice cream, cheese spreads, salad creams,
processed meats products, and pie fillings. It can also be used
to prepare a chocolate substitute, carob chocolate. A major use
of LBG, and guar, is in ice cream where it can act as a stabilizer
to prevent ice crystal growth at low temperatures.
Guar galactomannan has long been used as a food ingredi-
ent in India. Guar is a linear b14 mannan with a higher
proportion of galactose substituents than LBG, having a
galactose/mannose ratio of 38:62, and this is reflected in the
easier dispersion of guar when compared with LBG. Guar can
be dissolved at lower temperatures (20 ) as the extent of
unsubstituted regions of mannan chain is lower, reducing
opportunities for interchain hydrogen bonding that would
lead to aggregate formation and prevent hydration. The rheol-
ogy of guar is similar to that of LBG, pseudoplastic and decreas-
ing with temperature, with good pH stability. Gel formation is
only observed when other polysaccharides are added but the
ability of guar to participate in synergistic actions is lower than
that of LBG. Again, this is attributed to the reduced extent of
galactose-free regions of the mannan chain that could form
close hydrogen bonding with another polysaccharide.
Galactomannan from tara seeds has a galactose/mannose
ratio of 27:73, intermediate between those of LBG and guar.
The rheological properties are similar to those of LBG. Cassia
galactomannan has an average galactose/mannose ratio of
approximately 18:82 and can only be solubilized after boiling
when a high-viscosity solution can be obtained. The structure
of the gum is rather variable and fractions of different solubilit-
ies are obtained due to varying degrees of galactose
substitution.
Like the aforementioned tree seeds of the legume subfamily
Caesalpinioideae, the major grain legumes in the Papilionoi-
deae can also contain galactomannans as well as arabinoxylans.
The Papilionoideae include all the main bean crops of agricul-
ture including soya beans (Glycine max), Phaseolus spp., Vicia
faba, and Vigna spp. and may contain levels of total fiber from
20% to 50% with percentages of monosaccharides indicative of
arabinoxylans and galactomannans. One particular example is
fenugreek (Trigonella foenum-graecum), a legume from the Med-
iterranean region that contains high levels of a seed endosperm
galactomannan with a galactose mannose ratio approaching
1:1. Fenugreek galactomannan is cold-water-soluble forming
solutions with lower viscosity than the other galactomannans.
It has potential as an emulsifier and can show good ability to
stabilize oil/water interfaces. Soybeans contain a trace amount
of seed galactomannan with a galactose/mannose ratio
approaching 1:2, while lupin seeds (Lupinus albus) are unusual
in containing an unrelated reserve galactan (or arabinogalactan)
with a main chain of b14 linked galactose. Bean dietary fiber in
various forms is now available commercially as a by-product of
oil or protein extraction processes; however, the composition is
not well characterized and can contain high levels of cellulose
from the seed coat or husk. It is also marketed as bean bran or
hull flour for use as an ingredient to add dietary fiber to bakery
products.
Identifying NSPs
As new species are tested for potential as grain producers,
investigation for the presence of NSPs should be a component
in any study. It is relatively simple to ascertain the presence of
any unusual polysaccharides where these contain different
sugar units from those in known polymers. From the analysis
of the monosaccharide composition of a grain sample after
total acid hydrolysis, it will often be possible to infer the
presence of polysaccharides from the ratios and quantities of
monosaccharides present. Monosaccharide analysis is con-
ducted by HPLC or gas chromatography after generation of
volatile sugar derivatives. Where the polysaccharides differ
more subtly by variation in the linkages between similar
monosaccharides than the method of choice is usually meth-
ylation analysis, requiring the production of methylated sugar
derivatives reflecting the positions of glycosidic linkages that
can then be analyzed by GC/MS. Where soluble polysaccha-
rides are under investigation clear signals reflecting the posi-
tion of linkages can sometimes be obtained relatively rapidly
by NMR analysis.
The aforementioned analytic techniques will usually be
complemented by prior separation of polysaccharides present
into different fractions depending on their solubility in differ-
ent solvents. Using the water solubility/insolubility of a poly-
saccharide is a rapid way to simultaneously extract
polysaccharides from a sample and distinguish among those
present in the grain.
Conclusion
An enormous diversity of NSPs are potentially present in seeds.
The polysaccharides of the major cereal crop grains are well
known, but as the exploitation of grain resources expands to
include new and often diverse dicot plant resources, new NSPs
 CARBOHYDRATES | Grains other than Cereals: Non-starch Polysaccharides
will be encountered. Many of these new grains have not been
fully investigated for the composition of minor polysaccharides
present in the endosperm that may have a significant impact on
the properties of grain-derived products. In particular, grain
legumes as well-developed crops with high levels of NSP
would have potential for further investigation to establish the
composition of dietary fiber fractions and permit exploitation as
defined NSP sources. Similarly, there are opportunities for the
development of grain crops to obtain NSP resources that are
presently obtained from related species with much lower yields.
Eventually, we may see NSPs no longer being regarded as a
nuisance reducing the value of the grain but instead representing
the major economic product.
Exercises for Revision
What is a non-cereal grain? Give examples from different
plant groups.
Give examples of where NSP may be found in plant tissues.
What is the difference between NSP and dietary fiber?
What are the common chemical structures found in non-
cereal grain NSP?
Which non-cereal grain NSPs are exploited commercially
by the food industry?
What by-products from the agriculture industry may con-
tain NSP?
Exercises for Readers to Explore the Topic Further
Consider the various by-products available from legume crop
processing and suggest if any could be used to extract NSP.
Examine the geographic distribution of non-cereal grains
and cereal grains and consider to what extent they represent
complementary resources.
229
Using literature sources, try and find a new non-cereal grain
crop that is not currently exploited for NSP and predict
from DF content if it would be a useful source.
See also: Carbohydrates: Carbohydrate Metabolism; Cereals:
Chemistry and Physicochemistry of Non-starchy Polysaccharides;
Health Effects of b-Glucans Found in Cereals; Food Grains and
Well-being: Functional Foods: Dietary Fibers, Prebiotics, Probiotics,
and Synbiotics; Functional Foods: Overview; Processing of Grains:
b-Glucans: Measurement and Processing; The Basics: Grain:
Morphology of Internal Structure; The Legumes and
Pseudocereals: Grain Legumes and Their Dietary Impact: Overview;
The Oilseeds: Oilseeds: Overview.
Further Reading
Brett C and Waldron K (1996) Physiology and Biochemistry of Plant Cell Walls,
2nd edn. London: Chapman Hall.
Cui SW (2001) Polysaccharide Gums from Agricultural Products. Pennsylvania, PA:
Technomac.
Eliasson A-C (ed.) (1996) Carbohydrates in Food. New York: Marcel Dekker.
Harris PJ and Smith BG (2006) Plant cell walls and cell-wall polysaccharides:
Structures, properties and uses in food products. International Journal of Food
Science and Technology 41(s2): 129143.
Hedley CL (ed.) (2001) Carbohydrates in Grain Legume Seeds. Wallingford: CABI
Publishing.
Loewus FA and Tanner W (eds.) (1982) Plant Carbohydrates I. Encyclopedia of Plant
Physiology, vol. 13A. Berlin: Springer.
Nwokolo E and Smart J (eds.) (1996) Food and Feed from Legumes and Oilseeds.
London: Chapman & Hall.
Ouhida I, Perez JF, and Gasa J (2002) Soybean (Glycine max) cell wall composition and
availability to feed enzymes. Journal of Agricultural and Food Chemistry 50(7):
19331938.
Phillips GO and Williams PA (eds.) (2000) Handbook of Hydrocolloids. Florida: CRC
Press.
Tharanathan RN and Mahadevamma S (2003) Grain legumes A boon to human
nutrition. Trends in Food Science and Technology 14(12): 507518.
Tosh SM and Yada S (2010) Dietary fibres in pulse seeds and fractions:
Characterization, functional attributes, and applications. Food Research
International 43(2): 450460.
 Resistant Starch and Health
A Evans, Tate and Lyle, Hoffman Estates, IL, USA
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
What is resistant starch?
Resistant starch intake; measurement and consumption.
Effect of resistant starch intake on gut health.
Metabolic effects of resistant starch intake.
Learning Objectives
To get a basic understanding of what resistant starch is and
where it is found.
To learn about how its consumption has been linked to a
variety of potential beneficial health outcomes in the area
of gut health and various metabolic effects such as blood
glucose, insulin, and lipid levels.
What Is Resistant Starch?
Starch is the major source of carbohydrates in the human diet.
Starch is present in many different fruits, vegetables, roots, and
grains. Starch and starch derivatives are a nutritive, abundant,
and economical food source. Starch can be consumed unpro-
cessed in the form of raw fruits and vegetables or in the form of
more shelf-stable processed foods. Food starches contribute to
the characteristic viscosity, texture, mouthfeel, and consistency
of many food products.
In the human body, starch is digested by a-amylases. First,
salivary a-amylase in the mouth catalyzes the hydrolysis of
amylose to maltose, maltotriose, and maltotetraose and the
hydrolysis of amylopectin to the same products plus two
a-limit dextrins. The partially digested starch then passes into
the stomach. After some resident time in the low pH environ-
ment of the stomach, the partially hydrolyzed starch then
passes into the small intestine where it is neutralized. The
majority of hydrolysis of the starch is then accomplished by
pancreatic a-amylase that is secreted from the pancreatic duct
via a multiple attack mechanism. a-Amylases hydrolyze the
starch into small mono-, di-, and oligosaccharides that then
need to be further broken down to be absorbed as glucose. Two
other enzymes are necessary to convert the hydrolysis products
of the a-amylases into glucose, which can be actively trans-
ported across the small intestine membrane. These two
enzymes are the brush-border glucogenic enzymes maltase
glucoamylase and sucraseisomaltase. The resulting D-glucose
is then actively transported across the luminal membrane of
the small intestine and passes into the blood via the sodium
glucose cotransporter, which is located at the luminal surface
of enterocytes.
Several factors influence the rate and extent of starch diges-
tion. The main hydrolysis of starch is performed by the
a-amylases. Starch digestion by a-amylases requires a series of
230 Encyclopedia of Food Grains, Second Edition
steps. First, the enzymes diffuse into the starch matrix of the
food. In the second step, the enzymes bind to the substrate,
and finally, the enzymes cleave the a-1,4-glycosidic linkages of
the substrate (starch). The structure of the food matrix and the
structure of the starch itself influence the kinetics of the amy-
lase hydrolysis.
Resistant starch (RS) has been defined as the starch and
products of starch digestion that are not absorbed in the small
intestine of healthy individuals. Based on the source of the
enzyme resistance, RS has been classified into five different
types (Table 1).
In type 2, type 3, and type 5 RS, the enzyme resistance is due
to the physical structure of the starch molecules. Type 2 RS is
found in raw starch granules and the enzyme resistance is due
to the natural organization of the starch within the starch
granules. Significant amounts of type 2 RS can be found in
green banana, potato, and high-amylose maize.
In type 3 RS, the enzyme resistance is due to the physical
structure of starch chains that have undergone some type of
restructuring due to retrogradation or heat treatment. Different
factors, such as amylose/amylopectin ratio, chain length, lipid
content, and processing conditions, have been shown to influ-
ence the amount and quality of type 3 RS. Annealing and heat
moisture treatment are two types of heat treatments that are
often used to create RS. Both of these treatments, as well as
gelatinization, either fully or partially melt crystalline structure
present in the native starch granules. After heat treatment, linear
amylose molecules and linear regions of amylopectin molecules
can organize into a mix of amorphous and crystalline areas with
varying degrees of enzyme resistance. Formation of the crystal-
line structures usually takes place above the glass transition
temperature and below the melting temperature, and any com-
ponents present that influence the glass transition temperature
can therefore be expected to influence the formation (yield and
quality) of the formed type 3 RS. The amylose content of starch
has been positively correlated with RS yield and the formation
of type 3 RS is strongly related to the crystallization of amylose.
The amount of RS formed is also dependent on the water
content and temperature used during the heat treatment.
Water does act as a plasticizer in the system and a minimum
water content is necessary to achieve the chain mobility needed
to form crystalline structure resistant to enzyme digestion. At
high-starch concentrations, the starch chains interact more eas-
ily, leading to increased crystal and RS formation. The presence
of lipids has been shown to decrease the formation of type 3 RS
due to formation of amyloselipid complexes (type 5 RS).
The enzyme resistance in type 5 RS is due to the molecular
structure of amyloselipid complexes that can be either present in
the native starch or formed by controlled reactions using non-
granular starch and lipids to form the resistant amyloselipid
complex.
The enzyme resistance in type 4 RS is due to chemical mod-
ification of the starch. Chemical modification of starch creates
http://dx.doi.org/10.1016/B978-0-12-394437-5.00097-8

Table 1 Classification of resistant starch (RS)
RS
type Source of enzyme resistance Examples
1 Whole grains Seeds, grains
2 Raw starch granule structure Green banana, high
amylose, potato
3 Structures formed on Cooked pasta, cooked
reassociation or retrogradation potato
of starch after heat treatment
4 Chemical modifications of the Modified starch
starch
5 Starch lipid complex Amylose-containing starch
chain irregularities or branches in the starch chains. Cross-
linking of starch covalently links two starch chains together, in
effect creating a branch point on both chains. Chemical substi-
tution introduces a bulky side group to the starch chains. The
introduction of these chemical groups may create a steric hin-
drance to one or more human digestive enzymes.
RS Intake
Quantification of RS
To determine the effect of RS intake on health, it is important
to determine the amount of RS consumed as accurately as
possible. Ideally, RS consumption would be determined
in vivo by quantification of undigested starch in healthy indi-
viduals. This is, however, very difficult to do. Several in vivo
approaches have been used for measurement of RS. These
in vivo methods include the breath hydrogen test, direct collec-
tion of ileal effluent from ileostomy patients, and direct col-
lection of the ileal effluent from healthy subjects using a long
triple lumen tube.
The measurement of breath hydrogen to determine the
amount of RS in a diet is based on the concept of fermentation
of dietary fibers including RS. When RS enters the colon, it may
be fermented by the colon microbiota that leads to the pro-
duction of short-chain fatty acids (SCFAs) (acetate, butyrate,
and propionate) and gases like hydrogen (methane and carbon
dioxide). Most of the gases produced by colonic fermentation
are absorbed into the circulation, but about 10% is present in
the breath on exhalation. Since breath hydrogen is not specific
for only RS fermentation, but does indeed measure fermenta-
tion from any dietary fiber source, care has to be taken when
interpreting these results for RS content in a food/diet.
Direct collection of ileal effluents would be expected to give
more accurate results than breath hydrogen measurement
especially when different forms of dietary fibers and RS may
be present in a diet. Analysis of the ileal effluent allows for
quantification of undigested carbohydrate and potential iden-
tification of the different sources of dietary fiber within the
effluent. Most ileostomy models are, however, based on the
use of patients that had their colon removed due to health
issues and their digestive patterns may therefore not be repre-
sentative of a healthy population. Analysis of ileal effluent
from healthy individuals should provide the most accurate
CARBOHYDRATES | Resistant Starch and Health 231
data, but these measurements are difficult and intrusive and
can therefore not be done on large scales.
It is generally accepted that any in vitro method used to
measure RS should give values in line with those obtained
with ileostomy patients. In 1986, Berry developed a new
in vitro method by modification of a method previously devel-
oped by Englyst, Wiggins, and Cummings in 1982. This mod-
ified method was subsequently confirmed to give results in line
with healthy ileostomy subjects. In 1992, Englyst developed a
method that could measure rapidly digestible starch (RDS),
slowly digestible starch (SDS), and RS. This method employed
incubation of the samples at 37  C with pancreatic amylase
and amyloglucosidase and measures the amount of RS by
subtracting the sum of RDS and SDS from the total starch
present in the sample. Several new methods for the measure-
ment of RS were developed during the European Research
Program EURESTA. Even though many different methods
were developed during the 1990s for the measurement of RS,
none of these methods were successfully subjected to interla-
boratory evaluation to be adopted as an official AOAC method
of analysis. In 2002, McCleary and Managhan developed a new
RS method by modifying existing methods to obtain a method
that would yield robust, reproducible results that correlated
with results from ileostomy patients. This method underwent
interlaboratory evaluation and was later adopted as AOAC
Official Method 2002.02. An Integrated method for the mea-
surement of total dietary fiber was later developed and was
published in 2007. This method allows the accurate measure-
ment of all dietary fibers including RS. The enzymatic incuba-
tion step uses pancreatic a-amylase and more closely simulates
digestion in the human digestive tract and yields RS values in
line with those obtained with AOAC Official Method 2002.02
and with results from ileostomy patients. This method was
accepted as AOAC Method 2009.01.
Sources of RS and Intake Levels
Of all unprocessed foods, the green banana has the highest RS
content of between 47% and 57%. While flour can be prepared
from green bananas and incorporated into different foods as a
source of RS, it is important to keep in mind that food proces-
sing methods such as cooking may gelatinize the starch gran-
ules and destroy the structures responsible for the enzyme
resistance originally present and responsible for RS formation.
Raw potato starch is also high in RS, but since raw potato starch
is seldom consumed, it is not a good source of RS. However,
cooling of cooked potatoes or potato products can lead to
retrogradation of the starch and may lead to formation of
new RS (type 3) if the right conditions (heat, moisture, time,
and starch concentration) are met. The level of RS in different
legumes can vary widely and RS content in the legumes as
consumed depends on processing and legume variety.
Murphy et al. examined the RS intake in the United States
by comparing reported RS contents of different foods with
dietary recall records from participants in the 19992002
National Health and Nutrition Examination Survey. They
showed that the RS concentration varies greatly, even within
the same food category. This could be due to natural differ-
ences (cultivar), differences in food preparation methods, and
232 CARBOHYDRATES | Resistant Starch and Health
also differences in analytic methods used to determine the
amount of RS present. The RS content of different breads varied
between 0.1% in wheat rolls and 4.5% in pumpernickel bread.
Ready-to-eat breakfast levels contained between 0% (rice
cereal) and 6.2% (puffed wheat) of RS and cookies/crackers
had between 0.2% (oatmeal) and 2.8% (crackers and crisp
bread) of RS. The category of cooked cereal and pastas had
the widest range of RS contents between 0.4% (noodles and
chow mein) and 11.3% (oats and rolled uncooked). Within
the vegetable category, the lowest RS content was shown for
sweet corn (cooked/canned) and potatoes (slow-cooked) with
0.3% while the highest content was observed for fried potatoes
with 2.8%. In the legume category, RS content ranged from
0.6% for chickpeas (cooked/canned) to 4.2% for white beans
(cooked/canned).
Comparison of the RS content of different foods and the
food intake records showed that Americans on average con-
sumed an estimated 38 g of RS per day. In Europe, the intake
of RS has been estimated to be between 3 and 6 g per day while
the intake in developing countries has been estimated to be
between 30 and 40 g per day.
Health Benefits of RS
Gut Health
By definition, RS escapes digestion in the human small intes-
tine. RS therefore passes into the colon where like other dietary
fibers, it may be fermented. Fermentation of carbohydrates in
the colon leads to the production of SCFAs. SCFAs are the
metabolic products of anaerobic bacterial fermentation and
consist of butyrate, propionate, and acetate.
The SCFAs produced by bacterial fermentation in the
human gut are the preferred fuel for the cells lining the colon
(colonocytes) and have been shown to lower luminal pH,
increase the excretion of bile acids, and help prevent the devel-
opment of abnormal colonic cells. A lower pH has been asso-
ciated with protection against colorectal cancer and has been
shown to inhibit the conversion of primary to secondary bile
acids. Secondary bile acids are thought to promote tumors as
they are cytotoxic to colonic cells. It is therefore reasonable to
assume that RS may confer health benefits to gut health by
production of SCFA, which in turn have a positive effect on pH
and reduced production of secondary bile acids. A low pH has
also been found to suppress the growth of potentially patho-
genic organisms and reduce the absorption of toxic com-
pounds (such as ammonia).
Individual SCFAs have also been shown to have more spe-
cific actions in the colon. Most RS is readily fermented into
SCFA and many are a good source of butyrate. Butyrate is a
preferred energy source for colonocytes and helps drive the
uptake of electrolytes and water. Intake of butyrate producing
RS as part of oral rehydration solutions has been shown to
significantly reduce the recovery time for children with watery
diarrhea. Butyrate has also been shown to help maintain bar-
rier function in the gut by enhancing mucin production. The
barrier function is important to reduce bacterial translocation,
which may be a key element in the development of diseases
such as inflammatory bowel disease (IBD). Intake of RS has
been suggested to be beneficial for the reduction of IBD.
Butyrate has been shown to induce apoptosis, decreasing
the risk of cancer development. Animal studies have shown
that an increased supply of butyrate to the large bowel
decreases the induction of large bowel tumors in rats. How-
ever, despite the fact that the data strongly suggest the link
between butyrate and reduced risk of colonic cancer, studies
examining the link between RS intake and colonic cancer have
not been able to prove such a relationship. One reason for this
may be the importance of RS intake levels. Studies examining
the butyrate production from RS intake have shown significant
increase in fecal butyrate when 22 g of RS was consumed per
day, but no changes in butyrate or pH levels were observed
when 12.5 g of RS was consumed per day. The level of butyrate
produced may also depend on the type of RS consumed.
Another aspect of gut health often linked to dietary fibers is
laxation and fecal bulk. An increase of fecal bulk is thought to
be beneficial and is often reported as a potential health benefit
of different dietary fibers. Increases in fecal mass are important
for relief of constipation and prevention of diverticulosis and
other disorders such as hemorrhoids. The effect of RS on fecal
bulk has been studied and an increase of fecal output was
reported from different studies. Phillips et al. reported an
average increase in fecal output of 42% and also showed that
the fecal output increased with increased RS intake. This
increase fecal output could be due to increased water in the
fecal material, undigested and unfermented starch and dietary
fibers, or possible bacteria from the digestive tract. Increased
fecal nitrogen was observed in diets high in RS indicating either
an increase in bacterial mass or protein excretion.
Metabolic Effects
Several studies have shown that replacement of digestible car-
bohydrate with RS leads to a reduced blood glucose response.
This has formed the basis for approval of a European Food
Safety Authority (EFSA) claim for RS (EFSA, ID 681, April
2011). While this claim was granted, there was insufficient
evidence for the effects of RS when the glycemic load (amount
of digestible carbohydrate) remains constant. In 2012,
Robertson reviewed several studies on the effect of RS addition
to the diet when the available carbohydrate load was held
constant and noted a significant inconsistency between the
studies reviewed. While some found no effect of RS on glyc-
emia and significant reduction in insulin secretion, others
found the opposite, that is, a reduction in glycemia and an
elevation in postprandial insulinemia. The difference in these
results could be due to differences in the RS tested or simply be
due to the difficult relationship between glucose absorption,
clearance, and hepatic release, which makes accurate measure-
ments and conclusions difficult. Another important factor to
consider is the sources of RS used in different studies. RS can
come from foods such as beans, high-amylose corn starch, or
potatoes, and these foods all have different physiochemical
properties that can affect the response to RS ingestion. The fat
content of the diets is also important as fat is known to have a
significant impact on the glycemic response to a meal. A careful
review of studies with appropriate fat and protein levels
showed that RS consumption does indeed lead to a small
decrease in postprandial glycemia and a larger attenuation of
postprandial insulinemia. These acute changes in plasma

glucose and insulin concentrations caused by the addition of
RS to the diet could have important metabolic implications.
Both hyperglycemia and hyperinsulinemia are affected in the
development of insulin resistance. Increases in plasma glucose
concentrations have been linked to increased concentrations of
free fatty acids. Glucose uptake in the muscles and increased
glucose uptake in the adipose tissue have been linked to
hyperinsulinemia.
The potential chronic effects of RS intake on metabolism
and associated diseases are even more interesting than the
acute effects discussed earlier. There are animal data showing
a significant relationship of RS inclusion in the diet and
improved insulin sensitivity. Decreased insulin sensitivity, or
insulin resistance, has severe metabolic consequences as it is
strongly associated with several diseases known as metabolic
syndrome. Metabolic syndrome describes diseases such as type
2 diabetes, obesity, coronary heart disease, hypertension, and
dyslipidemia.
Even if RS only leads to reduction in blood glucose
response when digestible carbohydrates are replaced, such an
effect may have significant benefits to human health. Diabetes
affects about 8% of US population and is increasing globally.
Diabetes is characterized by hyperglycemia that can subse-
quently lead to systemic tissue toxicity. Risk factors for diabetes
include increased glucose response (both fasting and postpran-
dial), decreased insulin sensitivity, and obesity. These risk
factors may be reversible with lifestyle modifications that
have been shown to be effective in delaying the onset of type
2 diabetes. One lifestyle change could be to reduce the glyce-
mic load of the diet that could be achieved by replacing ordi-
nary starch in foods with RS.
RS has been shown to have significant effects of serum
triglyceride and cholesterol concentrations. Several animal
studies have shown significantly lower plasma cholesterol and
triglyceride concentrations when RS is included in the diet in
replacement of digestible starch. A decrease in fasting choles-
terol and triglycerides was observed when RS was consumed
chronically (514 weeks). The mechanism responsible for
these effects seems to be increased bile acid excretion caused
by consumption of RS, but some study showing reduced
plasma cholesterol and triglyceride concentrations even com-
pared to the bile acid sequestrant cholestyramine suggests that
the effects of RS extend beyond just bile acid secretion. The
effects of RS on cholesterol and blood triglyceride levels seem
to be mediated through a combination of enhancement of bile
acid secretion, a significant decrease in cholesterol absorption,
and an increase in hepatic LDL receptor expression. Both high
blood lipid and cholesterol levels are associated with coronary
heart disease and the effects of RS consumption may therefore
have important health ramifications.
The effects of RS consumption on satiety and weight man-
agement are not as clear. One major reason for the current
global obesity epidemic is the overconsumption of energy, and
therefore, strategies that can manage the energy intake may be
successful in reducing obesity. Dietary fiber is one way to
decrease the energy value of foods if it is used to replace
digestible carbohydrates that have a higher caloric value.
Some dietary fibers have also been linked to increased satiety
and lower BMI, but not all fibers have the same effects. RS is a
type of dietary fiber and it would therefore be reasonable to
CARBOHYDRATES | Resistant Starch and Health 233
expect some effect on satiety and/or total energy intake when
RS is used to replace digestible starch in the diet. Animal
studies have shown the production of satiety hormones
GLP-1 and PYY when RS was added to the diet, which suggests
an effect of RS on satiety. However, only a few human studies
have been conducted to examine the effect of RS on satiety and
the results were mixed. Some studies have shown an increase in
satiety after RS consumption, but others have shown little or
no effect. Satiety seems to be linked to changes in plasma
glucose concentrations and studies that showed a decrease in
plasma glucose/insulin response to a high RS meal also
showed an increase in satiety while those studies that showed
no change in plasma glucose also showed no difference in
satiety. Overall, there is indication that there is at least a weak
association between RS consumption and satiety. Some animal
studies have shown reduced energy intake when RS was added
to the diet. Aziz et al. showed that body weight was reduced by
40% in obese rats that were fed with a diet high in RS com-
pared with rats fed with a low-RS diet. While this study seems
to indicate a real positive effect of RS on energy intake and
potentially obesity management, the levels of RS used for this
study (23.4%) may not be realistic in a human diet. Long-term
studies in humans with realistic RS inclusion levels are
required to get a better understanding of the potential for RS
to help in the management and reduction of the current obe-
sity epidemic.
Summary/Conclusions
RS is a dietary fiber and can be used to replace digestible starch
in foods. Many animal studies have been conduced to examine
the effect of RS consumption. RS may be fermented in the
colon and the resulting SCFA production has been linked to a
number of health benefits such as reduced pH, which has been
associated with the suppression of growth of potentially path-
ogenic organisms and reduction of the absorption of toxic
compounds. Fermentation of RS has been shown to produce
relatively high levels of the SCFA butyrate that has been linked
to reduced risk of colon cancer. RS fermentation has also been
linked to benefits in the risk reduction of IBD by strengthening
of the gut barrier via increased production of mucin. RS has
been shown to increase fecal bulk, which is important for relief
of constipation and prevention of diverticulosis and other
disorders such as hemorrhoids.
An EFSA claim was granted in 2011 for reduced blood
glucose response when RS is used to replace digestible carbo-
hydrate in the diet (EFSA, ID 681, April 2011). The effect of RS
on glycemia and insulinemia is less clear, and studies have
shown that different effects are possible due to study designs
(e.g., amount of fat and protein in diets) and amount of RS
tested. Overall, it does seem that RS consumption does indeed
lead to a small decrease in postprandial glycemia and a more
significant attenuation of postprandial insulinemia that could
have important metabolic implications. Both hyperglycemia
and hyperinsulinemia are affected in the development of
insulin resistance, which in turn has been linked to several
diseases known as metabolic syndrome, for example, type 2
diabetes, obesity, coronary heart disease, hypertension, and
dyslipidemia.
234 CARBOHYDRATES | Resistant Starch and Health
RS has been shown to have significant effects on serum
triglyceride and cholesterol concentrations. The effects of RS
on cholesterol and blood triglyceride levels seem to be medi-
ated through a combination of enhancement of bile acid secre-
tion, a significant decrease in cholesterol absorption, and an
increase in hepatic LDL receptor expression. Both high blood
lipid and cholesterol levels are associated with coronary heart
disease and the effects of RS consumption may therefore have
important health ramifications.
The effects of RS consumption on satiety and weight man-
agement are not as clear. RS is a type of dietary fiber and it
would therefore be reasonable to expect some effect on satiety
and/or total energy intake when RS is used to replace digestible
starch in the diet. Only a few human studies have been con-
ducted to examine the effect of RS on satiety and the results
were mixed. Overall, there is indication that there is at least a
weak association between RS consumption and satiety, but
long-term studies in humans with realistic RS inclusion levels
are required to get a better understanding of the potential for
RS to help in the management and reduction of the current
obesity epidemic.
Exercises for Revision
What are the different types of resistant starch?
What is the link between resistant starch consumption and
colon cancer?
How does resistant starch affect insulin sensitivity and why
is this important?
Exercises for Readers to Explore the Topic Further
While many animal studies have been conducted examining
the relationship of resistant starch intake and various health
benefits, a more limited number of human clinical studies
have been conducted. The results of these clinical studies are
sometimes contradictory, and strong conclusions on the health
benefits of resistant starch are therefore difficult in some areas.
More human studies with careful study designs are needed in
the future.
See also: Carbohydrates: Carbohydrate Metabolism; Health Effects
of b-Glucans Found in Cereals; Food Grains and the Consumer:
Consumer Trends in Grain Consumption; Grains and Health:
Misinformation and Misconceptions; Grains and Health; Food Grains
and Well-being: Functional Foods: Overview; Grains Around the
World: Grain Production and Consumption: Overview; The Basics:
The Grains that Feed the World.
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 Health Effects of b-Glucans Found in Cereals
SM Tosh, Agriculture and Agri-Food Canada, Guelph, ON, Canada
S Shea Miller, Agriculture and Agri-Food Canada, Ottawa, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Mixed-linkage b-glucans are found almost exclusively in
members of the grass family.
Endosperm cell walls of oats and barley contain the highest
amounts of mixed-linkage b-glucans.
The b-glucan from oats, barley, rye, and wheat differs in size
and structure.
Solubility and viscosity are largely responsible for
b-glucans physiological effects.
Consumption of oat or barley products moderates glucose
response after a meal.
Over time, oat or barley reduces LDL cholesterol in hyper-
cholesterolemic individuals.
b-Glucan fermentation by colonic flora is associated with
improved gut health.
Learning Objectives
Mixed-linkage b-glucans are found almost exclusively in the
cell walls of members of the grass family, with the endo-
sperm cell walls of oats and barley containing the highest
amounts. The b-glucan from oats, barley, rye, and wheat
differs in molecular weight, as well as ratios of (1 ! 3) and
(1 ! 4) linkages. Differences in size and linkage ratios affect
the solubility and viscosity of b-glucan in solution which
are largely responsible for its physiological effects. Con-
sumption of products containing b-glucan from oats or
barley has been demonstrated to moderate the glucose
response after a meal. Consumption of products containing
b-glucan from oats or barley over time has been demon-
strated to decrease LDL cholesterol in hypercholesterolemic
individuals. Because of the b-linkages, b-glucan is not
digested by the enzymes of the upper digestive tract, but is
fermented by colonic flora to generate butyrate, propionate,
and acetate, short-chain fatty acids that are associated with
improved gut health.
Introduction
In Western society, chronic diseases including type 2 diabetes
and cardiovascular disease are becoming major concerns. It is
recognized that dietary factors contribute to the risk of devel-
oping these diseases and that obesity is also associated with
their incidence. Therefore, there is an interest in foods and
bioactive components of foods that could reduce the likeli-
hood of developing type 2 diabetes or cardiovascular disease.
In the early 1980s, a relationship between the consumption of
oats and oat bran and reduction of LDL cholesterol, a risk
factor for cardiovascular disease, was recognized. Subsequent
research identified the soluble fiber (1 ! 3)(1 ! 4)-b-D-glucan,
236 Encyclopedia of Food Grains, Second Edition
commonly called b-glucan, as the bioactive component. It also
reduces the blood glucose response following a meal. Signifi-
cant amounts of soluble b-glucan are also found in barley and
have been shown to have similar effects to oat b-glucan.
Distribution
b-Glucan is a mixed-linkage polysaccharide found almost
exclusively in the cell walls of members of the grass family or
Poaceae, which contains a large proportion of the worlds
economically important crops. Cereals, including oats and
barley, are members of the grass family, which belongs to the
subclass of monocots. Although b-glucans are present in the
cell walls of both vegetative and reproductive tissues, the high-
est amounts are located in the endosperm cell walls of oats and
barley, followed by rye and wheat, in decreasing order
(Figure 1). Only oats and barley are considered to have eco-
nomically (and physiologically) significant amounts. The
b-glucans in these four cereals differ in molecular weight and
in the ratios of oligosaccharides and linkages making up the
polymer and thus in solubility as well (Table 1). Rice and corn
do not contain significant amounts of mixed-linkage b-glucan,
while the dicots, including legumes, pseudocereals, and nuts,
do not have any.
The oat endosperm cell wall is approximately 85% b-glucan,
and within the oat endosperm, the greatest concentration of b-
glucan is found in the thick cell walls of the subaleurone layer
(Figure 1(a)). The walls of the central endosperm are much
thinner (Figure 1(b)), and there is a correspondingly lower
concentration of b-glucan in that area of the kernel. In barley,
the cell walls, which are also high in b-glucan ( 70%), are
typically a uniform thickness throughout the endosperm, so
the b-glucan is much more evenly distributed (Figure 1(c) and
1(d)). In rye, the predominant cell wall polysaccharide is arabi-
noxylan, with the b-glucan being found mostly in a layer on the
inside of the wall (Figure 1(e) and 1(f)). In the rye endosperm
cell walls shown in Figure 1(f), it can be seen that the staining of
b-glucans in the walls is less intense than that of the oats and
barley shown in (b) and (d), indicating a lower b-glucan content.
Wheat is unique, in that the b-glucan is found in highest con-
centration in the aleurone cell walls (Figure 1(g)), with smaller
amounts in the endosperm cell walls just inside the aleurone
layer (Figure 1(h)). In oats, barley, and rye, a thin layer of b-
glucan can also be detected on the inside of the aleurone cell
walls (e.g., Figure 1(a), 1(c), and 1(e)).
The main mixed-linkage b-glucan found in fungi, such as
mushrooms and yeast, is structurally different. It is a branched
polysaccharide with a b-(1 ! 3) backbone and b-(1 ! 6) link-
ages at the branch points. So, although they have similar
names, the biological functions of supplements containing
yeast and mushroom b-glucans are quite different.
http://dx.doi.org/10.1016/B978-0-12-394437-5.00096-6
 CARBOHYDRATES | Health Effects of b-Glucans Found in Cereals 237

Figure 1 Fluorescence micrographs of glycol methacrylate sections of cereals. (a), (c), (e) and (g) stained with Congo red to show b-glucan (red)
in the endosperm and aleurone cell walls, as distinguished from the autofluorescence of phenolics (blue) in the aleurone cell walls (oriented at
the top of each micrograph). (b), (d), (f), and (h) stained with calcofluor white to demonstrate b-glucan in the cell walls of the central endosperm
(bluewhite fluorescence). Bars in (a)(c), (d), and (g) represent 20 mm. Bars in (e), (f), and (h) represent 10 mm. (a) and (b) oat; (c) and
(d) barley; (e) and (f) rye; (g) and (h) wheat.

Table 1 Content and characteristics of mixed-linkage b-glucan found in major cereal grains

b-Glucan content (% of kernel) Molecular weight (106 g mol1) Trisaccharide/tetrasaccharide ratio

Oats 28% 2.53 1.72.4

Barley 210% 1.82.5 2.93.4
Rye 1.52.5% 11.3 2.73.6
Wheat 0.51.5% 0.30.7 3.74.8

Chemical Structure
Mixed-linkage b-glucan is a linear homopolymer of glucose,
similar in structure to cellulose ((1 ! 4)-b-D-glucan). However,
there are single b-(1 ! 3) linkages interspersed with the
b-(1 ! 4) linkages that make up about 30% of the linkages.
Because the b-(1 ! 3) linkages are not regularly spaced,
b-glucan does not form crystalline structures and fibrils like
cellulose; thus, it is much more soluble in water than cellulose
(Figure 2).
Of all the cereals, the b-glucan in oats tends to have the
longest molecules (highest molecular weight). In the native
grain (Table 1), the molecular weight is generally around
22.5 million g mol1 (12 000 glucose units). Approximately
40% of the b-glucan in uncooked oatmeal, oat bran, or oat
flour is soluble in water at 37  C in the presence of digestive
238 CARBOHYDRATES | Health Effects of b-Glucans Found in Cereals
Cellotetraosyl
unit
Glucose -(1 4) linkage
monomer
Figure 2 Schematic diagram depicting the chemical structure of a portion of a b-glucan molecule.
enzymes. Cooking or extrusion can increase the solubility. The
b-glucan in barley tends to be slightly smaller, about 1 million
g mol1, but has a higher ratio of cellotriosyl (trisaccharide) to
cellotetraosyl (tetrasaccharide) units than that in oats (tri-
saccharide/tetrasaccharide ratio ranges from 2.9 to 3.4 for bar-
ley, compared with 1.72.4 for oats). Wheat and rye contain
b-glucans of even lower molecular weight, and while rye has a
similar trisaccharide/tetrasaccharide ratio to barley, that of
wheat (3.74.8) is significantly higher than the other cereals
(Table 1). Higher trisaccharide/tetrasaccharide ratios and
lower molecular weight have been associated with increased
elasticity of b-glucan gels.
Health Benefits
Cholesterol Reduction
It has been well established that daily consumption of soluble
b-glucans can reduce total and LDL cholesterol. A recent meta-
analysis determined that persons with mildly elevated serum
cholesterol levels (3.05.0 mmol l1) who consumed at least
3 g of oat b-glucan per day experienced an average reduction in
LDL cholesterol of 0.240.29 mmol l1. Similar benefits have
been demonstrated with the consumption of barley b-glucan.
Barley and b-glucan isolated from barley lowered LDL-
cholesterol concentrations by 0.210.39 mmol l1 compared
with control. Health claims are permitted on food products
containing physiologically significant levels of b-glucans from
oats or barley in the United States, Canada, the European
Union, and Malaysia.
The viscosity generated by the b-glucan in the gut increases
excretion of cholesterol and bile acids. Bile acids are released
into the small intestine to emulsify fat in ingested food. Nor-
mally bile acids are reabsorbed lower down in the digestive
tract and recycled, but viscous fibers tend to physically entrap
them and carry them out of the body. To replenish what was
lost, in preparation for the next meal, the liver converts stored
cholesterol into bile acids, thus lowering the total cholesterol
levels.
Attenuation of Glycemic Response
Oat and barley foods have been shown to reduce blood glucose
concentrations after a meal compared to control foods and the
bioactive component has been demonstrated to be b-glucan
(Figure 3). Approximately 4 g of b-glucan per serving is
required to observe a significant physiological effect.
In the stomach, viscosity generated by b-glucan impedes the
mixing of food with gastric fluids and slows gastric emptying
rate. In the upper intestine, it hinders mixing of digestive
Cellotriosyl
unit
-(1 3) linkage
8
Whole wheat
Blood glucose conentration (mmol l1)
Oat bran
7
6
5
4
3
0 20 40 60 80 100 120
Time (min)
Figure 3 Glycemic response to oat bran muffin containing 4 g of
b-glucan and whole-wheat muffin containing <0.1 g of b-glucan. After
consumption of the oat bran muffin, blood glucose levels increased
more slowly, peaked at a lower level, and returned to baseline
more slowly than after consumption of the whole-wheat muffin.
enzymes with food, inhibiting the release of glucose from
starch. Intact cell walls in cooked whole kernels (including
pearled and pot barley), steel-cut oat, and milled flakes also
protect starch from digestive enzymes.
Sugars, inherent in a meal or released from starch, take
longer to reach the intestinal wall in a high viscosity mass
than in low viscosity environment because of the slow mixing
and diffusion rates. Therefore, glucose enters the bloodstream
at a slower rate and over a longer section of the small intestine.
The protracted release of glucose into the bloodstream results
in a beneficial reduced insulin response; repeated spikes in
insulin concentrations increase the risk of developing glucose
intolerance that in turn may lead to type 2 diabetes.
Increased Satiety and Weight Management
Increased satiety and delayed hunger have been observed after
a meal containing oat or barley foods compared with foods
containing refined wheat flour. Although the mechanism is not
well established, the effect appears to be related to slower
gastric emptying and digestion rate, which is also responsible
for reduced glycemic response. In addition, changes in gut
hormone responses have been observed following consump-
tion of oat and barley foods. Despite changes in satiety, results
 CARBOHYDRATES | Health Effects of b-Glucans Found in Cereals 239

A A
* *

A
A
*
(a) (b) (c)

Figure 4 In vitro digestion and fermentation of oat bran. (a) Suspension of oat bran in buffer, stained with calcofluor to demonstrate b-glucan. Note
sheets of aleurone cells (A) and endosperm cell walls (*). (b) Suspension of oat bran after in vitro digestion with a-amylase and proteases to mimic
digestive enzymes, stained with calcofluor. Note that aleurone cell wall sheets (A) are unchanged by digestive enzymes but that the b-glucan in the
endosperm cell walls has been solubilized and appears as an amorphous blue mass (*), although some endosperm cell wall structure is visible (arrow).
(c) Suspension of oat bran/feces after in vitro fermentation, stained with calcofluor. The b-glucan has been completely fermented by the fecal flora.
Some autofluorescence of phenolics is visible in vascular elements from other plant-based foods (arrow), as well as in small fragments of blue cell walls
embedded in the larger fecal particle. Bars in (a)(c) represent 100 mm.

for studies measuring changes at a subsequent meal have been
mixed. Some studies observed a significant drop in calories
consumed while others did not.
Fermentation and Laxation
Oat and barley fiber are not broken down by enzymes in the
upper intestine and arrive in the colon intact. Intact oat bran
contains particles of aleurone and endosperm cells, which are
easily seen with the fluorescence microscope due to the auto-
fluorescence of phenolics in the aleurone layer and the blue
fluorescence of b-glucans stained with calcofluor (Figure 4(a)).
After an in vitro digestion with a-amylase and proteases meant
to mimic conditions in the upper GI tract, the b-glucans have
been mostly solubilized, as demonstrated by the loss of dis-
cernible cell wall structures and the appearance of amorphous
blue-staining material in Figure 4(b), but are still present in the
digesta. Because of its solubility, b-glucan is susceptible to
enzymes produced by gut microbiota and rapidly fermented,
as demonstrated by the loss of calcofluor-stained material in the
digesta (Figure 4(c)).
During fermentation of b-glucan, gut bacteria produce
increased levels of butyrate, propionate, and acetate, the
short-chain fatty acids associated with improved gut health
(and possibly satiety as well, in the case of acetate).
Oat bran improves laxation mainly because of the insoluble
fiber portion. Wheat bran is more effective at promoting laxa-
tion because of its high insoluble fiber content.
Exercises for Revision
Oat and barley b-glucans are not absorbed into the blood-
stream. How are they able to influence concentrations of
glucose and cholesterol in the blood?
b-Glucan is a dietary fiber. What is the definition of dietary
fiber? What functions do soluble and insoluble dietary fib-
ers play in maintaining health?
If wheat contains b-glucan, why does it not reduce
cholesterol?
Exercises for Readers to Explore the Topic Further
How do different types of food processing affect the effec-
tiveness of b-glucan? Does it matter whether oats and barley
are consumed in extruded breakfast cereals, cooked cereals,
breads, soups, snack bars, desserts, or beverages?
How can consumers be encouraged to increase their intake of
oat and barley foods to take advantage of their health benefits?
Oat and barley foods have been reported to increase and
extend satiety after a meal? Is the satiety effect related to b-
glucan or glycemic response modulation?
See also: Carbohydrates: Carbohydrate Metabolism; Cereals:
Chemistry and Physicochemistry of Non-starchy Polysaccharides;
Glycemic Index, Glycemic Load, and Their Health Benefits; Food
Grains and the Consumer: Grains and Health; Food Grains and
Well-being: Functional Foods: Overview; Processing of Grains:
b-Glucans: Measurement and Processing; The Cereal Grains:
Oats: Overview.

Where are b-glucans mainly found? Name the plant family.
Which food ingredients are the main sources?
Draw a schematic diagram of a b-glucan molecule includ-
ing two tetrasaccharide and two trisaccharide segments.
Identify the 1 ! 3 and 1 ! 4 linkages.
What are the linkage differences between cereal b-glucans,
cellulose, and starch? What are the implications for struc-
ture and physiological effect?
What four health benefits have been attributed to oat and
barley b-glucans?
Further Reading
Abdel-Aal EM and Wood PJ (eds.) (2005) Specialty Grains for Food and Feed. St. Paul,
MN: AACC International.
AbuMweis SS, Jew S, and Ames NP (2010) b-Glucan from barley and its lipid-lowering
capacity: A meta-analysis of randomized, controlled trials. European Journal of
Clinical Nutrition 64(12): 14721480.
Aman P (2006) Cholesterol-lowering effects of barley dietary fibre in humans: Scientific
support for a generic health claim. Scandinavian Journal of Food and Nutrition
50(4): 173176.
Anderson JW (1987) Dietary fiber, lipids and atherosclerosis. American Journal of
Cardiology 60: 17G22G.
240 CARBOHYDRATES | Health Effects of b-Glucans Found in Cereals
Braaten JT, Wood PJ, Scott FW, et al. (1994) Oat b-glucan reduces blood cholesterol
concentration in hypercholesterolemic subjects. European Journal of Clinical
Nutrition 48: 465474.
Chu Y-F (ed.) (2013) Oat: Nutrition and Technology. Chichester: Wiley.
El Khoury D, Cuda C, Luhovyy BL, and Anderson GH (2012) Beta glucan: Health
benefits in obesity and metabolic syndrome. Journal of Nutrition and Metabolism,
Article number 851362.
Kale M, Hamaker B, and Bordenave N (2013) Oat b-glucans: Physicochemistry and
nutritional properties. In: Chu Y-F (ed.) Oat: Nutrition and Technology. Chichester: Wiley.
Newman RK and Newman CW (2008) Barley for Food and Health: Science, Technology,
and Products. St. Paul, MN: AACC International.
Othman RA, Moghadasian MH, and Jones PJH (2011) Cholesterol-lowering effect of oat
b-glucan. Nutrition Reviews 69(6): 299309.
Shewry PR and Ullrich SE (eds.) (2014) Barley: Chemistry and Technology, 2nd edn.
St. Paul, MN: AACC International.
Tosh SM (2013) Review of human studies investigating the post-prandial blood-
glucose lowering ability of oat and barley food products. European Journal of
Clinical Nutrition 67: 310317.
Truswell AS (2002) Cereal grains and coronary heart disease. European Journal of
Clinical Nutrition 56: 114.
Webster FH and Wood PJ (eds.) (2011) Oats: Chemistry and Technology, 2nd edn. St.
Paul, MN: AACC International.
Whitehead A, Beck EJ, Tosh SM, and Wolever TMS (2014) Cholesterol-lowering effects
of oat beta-glucan: A meta-analysis of randomized controlled trials. The American
Journal of Clinical Nutrition 100: 14131421.
Wood PJ (2010) Oat and rye b-glucan: Properties and function. Cereal Chemistry
87(4): 315330.
Wood PJ (2011) Oat b-glucan: Properties and function. In: Webster FH and Wood PJ
(eds.) Oats: Chemistry and Technology, 2nd edn., pp. 219254. St. Paul, MN:
AACC International.
Wood PJ, Arrigoni E, Shea Miller S, and Amado R (2002) Fermentability of oat and
wheat fractions enriched in b-glucan using human fecal inoculation. Cereal
Chemistry 79: 445454.
Wursch P and Pi-Sunyer FX (1997) The role of viscous soluble fiber in the metabolic
control of diabetes: A review with special emphasis on cereals rich in b-glucan.
Diabetes Care 20: 17741780.
 Glycemic Index, Glycemic Load, and Their Health Benefits
D Dan Ramdath, Agriculture and Agri-Food Canada, Guelph, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Carbohydrates are required as part of a healthy diet, but
they vary in the rate at which they increase blood glucose
following ingestion.
Glycemic index (GI) is a numeric physiological basis of
classifying carbohydrate foods initially used to improve
blood glucose control in persons with diabetes.
Glycemic load is the product of the carbohydrate content
per serving of food and its GI.
Measurement of GI requires experienced users and a
method that is accurate and valid.
Glycemic response is the blood glucose response following
consumption of digestible carbohydrate and is measured as
the area under the blood glucose response curve.
Several food characteristics may affect derivation of GI
values.
Regular consumption of diets that are low in GI and GL is
associated with lower risk for type 2 diabetes (T2DM) and
cardiovascular disease (CVD).
Learning Objectives
To describe the different types of carbohydrates in the
human diet and their effects on blood glucose.
To describe the concept of glycemic index and glycemic
load and their role in management of diabetes and health
promotion.
To understand the derivation, required methodology, and
classification of GI.
To calculate the GI and GL of a mixed meal.
To understand the food characteristics that may affect der-
ivation of GI values.
To outline the difference between GI, GL, and glycemic
response.
To examine human studies on the role that GI and GL
might play in reducing risk for T2DM and CVD, and pro-
moting satiety and weight loss.
Introduction
Carbohydrates provide approximately 5060% of energy
intake in the normal human diet, and this requirement is met
through regular consumption of starchy foods such as cereals,
fruits, and vegetables. Following consumption, carbohydrates
are progressively broken down by digestive enzymes in the
gastrointestinal tract where degraded starch structures give
rise to glucose, which is then actively transported to various
tissues to meet cellular energy requirements. About four
decades ago, it was recognized that carbohydrates differ in
the rate at which they are digested and consequently vary in
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00098-X
their effects on blood glucose levels. In 1981, the glycemic (or
glycaemic) index (GI) was proposed as a numeric physiologi-
cal basis of classifying carbohydrate foods in order to improve
blood glucose control in persons with diabetes. Studies that
preceded this concept demonstrated that different carbohy-
drate foods raised blood glucose by varying amounts, which
could not be predicted from chemical analysis. Indeed, differ-
ences were measured among foods belonging to the same food
group. It is now widely accepted that the blood glucose raising
potential of carbohydrate-rich foods can be ranked in a stan-
dardized matter using GI, and this feature can be used as an
adjunct in the dietary management of persons with diabetes.
In 1997, a joint Food and Agriculture Organization/World
Health Organization Expert Consultation defined GI as the
incremental area under the blood glucose response curve
(AUC) elicited by a 50 g available carbohydrate (AC) portion
of a test food expressed as a percentage of the response elicited
by the same amount of AC from a standard food (either white
bread or glucose) taken by the same person. GI is therefore a
measure of how much a standard quantity of a food raises
blood glucose compared with a standard quantity of glucose
or white bread, with a value of 100 having been assigned to
glucose. In practical terms, GI is derived from repeated testing
of a carbohydrate food in at least ten research participants, with
the reference food tested at on
2 occasions (see eqn [1]).
Although the significance of GI in managing persons with
diabetes is widely accepted, there have been numerous debates
on the derivation of GI, including the accuracy and precision of
the method and its universal application to public health
promotion campaigns to guide healthy food choices:
Glycemic index food AUCfood =mean AUCreference  100 [1]
The derivation of GI is based on the amount of available (or
preferably glycemic) carbohydrates present in a carbohydrate-
rich food. GI compares equal quantities of carbohydrate and
provides a measure of carbohydrate quality but not quantity. It
has been argued that GI is a property of carbohydrate-rich
foods, which is independent of age, gender, body mass index,
ethnicity, and glucose tolerance status but may be influenced
by factors such as food variety, cultivars, processing,
preparation, and storage effects, among others. Using stan-
dardized methodology, the GI of carbohydrate-rich foods can
be assessed with a standard deviation of about 9, and the
resulting values can be classified (with glucose as the standard)
as high (GI
70), medium (GI 5669), or low (GI  55). See
Table 1 for a sample listing of GI values of different foods.
Diets containing high amounts of refined carbohydrates
have been shown to increase plasma levels of triglyceride and
reduce high-density lipoprotein. Assessing this type of relation-
ship in human studies required an understanding of carbohy-
drate quality (i.e., GI) and the quantity eaten in a meal or
during the course of the day. The concept of glycemic load
(GL) was therefore introduced in 1997 by researchers at
241
242 CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits

Table 1 Glycemic index values for various foods (glucose 100)

Fruits Root vegetables Legumes

Apple, Golden Delicious 39  3 French fries, baked (OreIda) 64  6 Baked beans in tomato sauce 40  5
Banana (Canada) 62  9 Mashed potato, instant (Canada) 80  13 Chickpeas, canned 38  3
Cherries, dark, raw 63  6 Parsnips, peeled, boiled 52  5 Chickpea hummus 22  5
Kiwi 58  7 Potato, Irish, peeled, baked 83  6 Kidney beans, dried, soaked overnight, boiled 45 min 51  5
Mango 51  3 Potato, Irish, boiled 59  4 Lentils, brown, canned 42  5
Orange (South Africa) 33  6 Sweet potato, peeled, boiled 46  5 Lentils, red, dried, boiled 21  7
Papaya 56  6 Taro (China) 48  5 Mung bean, germinated 25  4
Pineapple 66  7 White yam, boiled (Jamaica) 79  4 Soya beans, canned (Canada) 14  2

Dairy High-carbohydrate foods Snacks

Ice cream, vanilla (Sara Lee) 38  3 All-Bran (Canada) 51  5 Chips (Doritos original) 42  4
Milk, full-fat (Canada) 34  6 Cornflakes (Canada) 80  6 Chips, potato, plain, salted 51  7
Milk, skim (Canada) 32  5 Rice, Basmati, boiled 8 min 69  6 Chocolate, dark, DoveW 23  3
Soy milk, VitasoyW organic 43  3 Rice, brown 66  5 Chocolate, milk, DoveW 45  8
Yogurt, white peach (Finest) 34  7 Rice, sticky 87  2 Jelly beans (Allens) 80  8
Yogurt, Greek style, plain 12  4 Rolled oats, cooked 20 min 49  8 Mars BarW 62  8
Yogurt, frozen, vanilla Blue Menu 46  5 Spaghetti, white, durum wheat 58  7 Mixed nuts, roasted, salted 24  10
Yogurt, natural, fat-free 19  5 Wonder, white bread 72  4 Sports bar, Clif barW, Brownie 57  6

Low GI < 55; medium GI 5569; high GI  70.

Source: Atkinson FS, Foster-Powell K, and Brand-Miller JC (2008) International tables of glycemic index and glycemic load values: 2008. Diabetes Care, 31, 22812283.

Harvard University to quantify the overall glycemic effect of a
portion of food (see eqn [2]). GL is defined as the product of
the carbohydrate content per serving of food and its GI. As
such, dietary intakes of carbohydrates can be weighted to pro-
vide the GL, which is a summative value of the quantity of a
carbohydrate-rich food ingested and the quality of the food
and allows for a better understanding of how different-sized
portions of carbohydrate foods compare in terms of their
blood glucose raising effect. The higher the GL, the greater
the expected elevation in blood glucose and in the insulino-
genic effect of the food. Foods can be classified as having GLs
that are low (GL  10), medium (10 < GL < 20), or high
(GL
20).
GIfood  available carbohydrate g=serving
Glycemic loadfood
100
[2]
GI Derivation and Methodology
The method for derivation of GI has evolved considerably over
the last two decades with many improvements in the tech-
niques and prerequisites for testing. In an attempt to standard-
ize and improve the precision of the method used to determine
GI, two multilaboratory studies were undertaken with each lab
analyzing the same foods. These studies concluded that varia-
tion in the GI values obtained by established laboratories
could be accounted for by differences in data analysis and
within-subject variation of glycemic responses. Several
improvements in the methodology were suggested, including
duplicate measurements of fasting blood glucose, use of capil-
lary blood sample, and pretest restrictions on exercise and
alcohol consumption. Many of these improvements have
been included in the current International Organization for
Standardization methodology (ISO 26642:2010) for deter-
mining the GI of foods. This well-defined set of guidelines
include mandatory procedures, recommendations for charac-
teristics of study participants and pretesting preparation, and
internal validity checks, such as the use of a method to measure
blood glucose with an analytic variation of <3% and exclusion
of outliers.
Typically, the ISO GI method calls for a minimum of ten
healthy research participants who, following an overnight fast,
are randomly fed a 50 g AC portion of the test or reference food
(white bread or glucose) using a randomized block design.
Before consumption of the test meal, two fasting blood sam-
ples are obtained via the finger prick method, and additional
blood samples are collected at 15, 30, 45, 60, 90, and 120 min
after the first bite is taken. Persons with type 2 diabetes
(T2DM) or glucose intolerance can be included as research
participants, in which case additional blood samples are
taken at 150 and 180 min. Finger prick capillary blood is
preferred because (i) it is easier to obtain, (ii) the rise in
blood glucose is greater than in venous plasma, and (iii) results
for capillary blood glucose are less variable than those for
venous plasma. As such, differences in the blood glucose
response between foods are larger and easier to detect statisti-
cally using capillary blood glucose. Changes in the blood
glucose concentrations above fasting are calculated as the
incremental area under the glucose response curve using the
trapezoid rule. The reference food is eaten by each research
participant on at least two separate occasions to derive a mean
AUC for that individual. The AUC of each participant after each
test food is expressed as a percentage of the mean AUC elicited
by the reference food in the same participant. Next, the mean
of these values for all participants is calculated as the food GI. If
white bread is used as the reference food, the GI values are
multiplied by 0.71 to convert them to the glucose scale
 CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits 243

(i.e., the GI of glucose 100). The concept of GI testing and before the existence of a standardized method for GI and
derivation is illustrated in Figure 1. Given the care required to that in some instances, the GI values of the same food item
produce accurate and valid GI value, it is advisable that new have relatively large variation. Some of this variation could be
investigators adhere closely to the ISO method and perform a attributed to foods being tested in 10 persons; however,
minimum of three tests on the reference food. differences in GI values of the same food category may also
From the protocol outlined in the preceding text, it be explained by differences in processing, in determining
becomes clear that GI is not a simple ratio of two values, but the digestible carbohydrate content of the test foods, in GI
rather a summation of 3 or 4 values from each volunteer and testing, inherent botanical differences and growing conditions,
may be expressed mathematically as etc. Indeed, derivation of GI values is affected by many food
Xx1 factors (see Table 2), and this has been the focus of many
x
100  Fx =G disagreements on the relevance and application of GI to
GI n
[3]
n human health.
where Fx is the incremental blood glucose response (AUC) in
participant x elicited by 50 g AC from the test food and Gx is the
mean AUC in participant x elicited by 50 g glucose (or the Glycemic Carbohydrates
equivalent white bread) tested on two or three separate occa-
sions. In the hands of experienced investigators, this approach An imperative of GI testing is that both the reference foods and
with 12 healthy adult volunteers usually provides a power of test foods must contain the same amount of available or glyce-
80% to detect a 22% difference in AUC. Further, among 28 mic carbohydrate; typically 50 or 25 g portions are consumed by
centers participating in an interlaboratory study on GI meth- participants. The method used for determining available
odology, the between-laboratory standard deviation of the GI
values was 9.
Table 2 Food characteristics that may affect derivation of GI values
In order to reduce unnecessary repetition in the testing of
individual foods and to promote wider research and applica- Food factor Effect on GI
tion, an international table of GI and GL values was compiled
from the scientific literature in 1995 and updated in 2002 and Presence of monosaccharides Formulations with glucose will have
2008 to contain 2487 separate entries from 205 published Glucose; fructose; galactose higher GI than those containing
articles. Further, GI values derived from studies of 5 persons sucrose or high-fructose corn
syrup
are listed separately, and caution is given in using these values.
This resource, a sample of which is shown in Table 1, has been Type of starch Amylopectin has branched structure
instrumental in improving the quality of research examining Relative abundance of containing a1-6 linkages, which is
the relation between GI, GL, and health. It has been used by amylose and amylopectin digested faster and has higher GI
researchers in many human cohort and interventional studies Storage Many fruits and vegetables undergo
and in supporting dietary guidelines for high-fiber, low-GI Ripening ripening during which starch
foods for individuals with diabetes as a means of improving Moisture content decreases and sugars
postprandial glycemia and weight control. However, others Temperature increase
have been more reluctant to putting advice about GI values Retrogradation Moisture influences retrogradation
into practice over concerns about potentially adverse effects on and formation of resistant starch
Cold storage after cooking can
food choice and fat intake. Limitations in the use of the
promote recrystallization of the
international GI table include the fact that it was compiled starch chains
Processing Different cultivars often have
Starch particle size different starch structures
Cell wall integrity Large starch granules have larger
surface areas and higher GI
10
50 g glucose Disruption of cell wall during
Plasma glucose (mmol l1)

standard processing
Cooking Differences in the rate of
8 50 g AC Same product with different carbohydrate digestion and hence
Test food ingredients the GI value
Ingredients processed Degree of starch gelatinization and
6 differently consequently the GI values
Recipe variation; flour used,
moisture content, cooking
time
4
Other food components Soluble viscous fiber associated
Dietary fiber and resistant with lower GI
0 30 60 90 120 starch Presence of a-glucosidase inhibitors
Time (min) Antinutrients Delayed gastric emptying

Figure 1 Derivation of Glycemic Index

Organic acids
244 CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits

carbohydrate is therefore critical. In fact, there have been sugges-
tions that glucose should be used as the only reference food
because of the ease of measuring 50 g of glucose. AC is defined
as the fraction of carbohydrate that can be digested by human
enzymes, is absorbed, and enters into intermediary metabolism.
It does not include dietary fiber, defined as intrinsic plant cell
wall polysaccharides, which can be a source of energy only after
colonic fermentation in the large intestine. It has been recom-
mended that the term AC be replaced by glycemic
carbohydrate. Using this terminology, glycemic carbohydrates
will include most monosaccharides and disaccharides, some
oligosaccharides (maltodextrins), and rapidly digested starches.
Slowly digested starches, which are digested less rapidly to release
glucose, are also considered to be glycemic carbohydrates. In
contrast, nonglycemic carbohydrates include oligosaccharides,
nonstarch polysaccharides (defined as macromolecules consist-
ing of a large number of monosaccharide residues joined to each
other by glycosidic linkages), and resistant starch. Resistant
starch (RS) is defined as the sum of starch and products of starch
digestion (such as maltose, maltotriose, and a-limit dextrins)
that are not absorbed in the small bowel and can be classified
into four types: RS I, physically inaccessible starch mostly present
in whole grains; RS II, RS granules (B-type starches); RS III,
retrograded starch (after food processing); and RS IV, modified
starches.
It is possible that earlier GI studies relied on generic food-
composition tables, data from food manufacturers, or even
food labels, which are insufficient to obtain an accurate mea-
surement of glycemic carbohydrate. Accurate laboratory mea-
surement of glycemic carbohydrate should be undertaken prior
to embarking of GI testing. Typically, this involves analysis of
moisture, fat, ash, and protein from which total carbohydrate
P
is obtained as 100  ash protein moisture fat. In
addition, measurement of total dietary fiber is required.
Glycemic carbohydrate is then calculated as [total carbohy-
drate by difference  total dietary fiber]. This approach does
not take into account resistant starch, which is difficult to
measure routinely. There are suggestions that it may be more
accurate to quantitate glycemic carbohydrates directly, rather
than by difference; however, this measurement is only avail-
able in specialized laboratories.
GI Versus Glycemic Response
One of the central issues driving the debate on the derivation
and application of GI to human health is possibly an appreci-
ation of the difference between glycemic response and GI. As
shown in Figure 1 and outlined in eqn [3], a test or reference
food will produce an individual glycemic response, defined as
the blood glucose response following consumption of digest-
ible carbohydrate and measured as the AUC. As outlined
before, the food GI obtained in an individual is derived by
dividing the test food AUC by the mean glycemic response
obtained from 2 or 3 tests of the reference food in the same
individual. The food GI is then taken to be the average of the GI
obtained in at least ten individuals. This approach reduces
variation and provides a more precise estimate of the true
mean. Studies have shown that when healthy or diabetic vol-
unteers repeatedly consume carbohydrate foods (e.g.,
spaghetti, rice, and bread) on different occasions, significant
variation in their glycemic response is observed, but mean GI
values remain similar. Figure 2 probably best illustrates the
difference between glycemic response and GI and the effect of
individual variation on the derivation of these indices. Follow-
ing the consumption of 50 g AC portions of five different foods
(sucrose, instant potato, bread, rice, and barley) by persons
with varying levels of glucose tolerance, significant differences
in their blood glucose response were observed, but the
between-individual variation in GI of the foods studied
remained relatively constant. As such, it may be concluded
that GI is a property of the food, whereas the glycemic response
is a variable individual characteristic.
The ecological validity of dietary GI and GL has been
assessed in a randomized trial of lifestyle modification in

Partitioning of variance
12 AUC
1200 Within-subjects
Variance (106)

Between-subjects GI
Glycaemic index (%)

Food
AUC (mmol min l1)

4
800
0
AUC GI
120
400 80
40
0 0
1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31
Subject ID number
Figure 2 Comparison of variance associated with AUC and GI in persons with varying glucose tolerance.
Incremental AUC and estimates of GI in 31 study participants (9 normal, 12 hyperinsulinemic, and 10 with type 2 diabetes) elicited by 50 g avCHO
portions of five different carbohydrate foods (instant potato, white bread, spaghetti, barley, and sucrose); values are mean  s.d. for the five test meals in
each participant. AUC differs significantly among participants, but GI does not. Inset: the amounts of variance in AUC and GI due to the different foods,
the different study participants (i.e., between individuals), and day-to-day variation (within participants) from repeated-measures analysis of variance.
Expressing results as GI reduced variation between participants by 95%. Figure re-drawn using data derived from Lan-Pidhainy X and Wolever TMS
(2011) Are the glycemic and insulinemic index values of carbohydrate foods similar in healthy control, hyperinsulinemic and type 2 diabetic patients?
European Journal of Clinical Nutrition, 65, 727734.
 CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits 245

free-living overweight and obese individuals with T2DM, similar names (e.g., bread, cookies, and cakes) may have dif-
using continuous glucose monitoring to measure the glyce- ferent ingredients (e.g., type of flour used) or may have used
mic response to foods. In this study, participants consumed different processing methods (e.g., different cooking times
foods in self-selected amounts, at self-selected times, and in resulting in different moisture contents), which could influ-
self-selected combinations. Participants kept 3-day dietary ence the degree of starch gelatinization and rate of digestion
diaries from which intakes of energy, fat, protein, carbohy- and ultimately the GI value.
drates, sugar, and fiber were obtained and GI and GL calcu- Starch type and structure and relative abundance of amylose/
lated. After controlling for energy and carbohydrate intakes, amylopectin in different varieties of rice and potato can influ-
dietary GI accounted for 1018% of the variance in the ence their GI. Amylose is a form of starch with a long, straight
glycemic response indices assessed. Further, when dietary chain structure, whereas amylopectin is highly branched and
intake was adjusted for energy, GL and GI were significantly allows easier access by the digestive enzymes and is therefore
correlated with the glycemic response. The authors concluded associated with higher GIs. Various factors affect the amylose
that GI was the strongest and most consistent independent and amylopectin content of foods; many of these influence
predictor of glycemic response and variability. Less invasive retrogradation and resistant starch formation and include mois-
studies have also shown that the predominant determinants ture, cooking and storage temperature, reheating, and ripening.
of the glycemic response to a mixed meal are the amount of These factors are discussed in detail in other articles. Many fruits
carbohydrate and meal GI, which explain 90% of the varia-
tion in mean AUC. In fact, the glycemic response is depen-
dent on the relative amounts of carbohydrates and the
amounts and types of fat and protein in the mixed meal. Table 4 Glycemic index of a sample of relatively unprocessed foods
For example, it has been shown that addition of fat to a from different countries
potato test meal reduces the glycemic response by about
Food item Country GI (mean  SEM if available)
50%. Other factors affecting glycemic response include nature
of monosaccharide components, starch structure and relative White flour The United Kingdom 59  11
abundance of amylose and amylopectin, degree of ripening of Canada 69  5
fruits and vegetables, and cooking and processing. It is there- South Africa 71  7
fore reasonable to conclude that eating a food as part of a Italy 89  12
mixed meal alters the glycemic response, but not its GI; this Rice India 43
China 83  1
differentiation therefore allows for the calculation of the GI
Canada 72  9
of a meal containing several carbohydrate foods as outlined
Banana Australia 47  5
in Table 3. Canada 62  9
South Africa 70  5
Lentils Canada 29  3
Australia 37  3
Effect of Food Processing on GI/GL The United Kingdom 21  7
Soya beans Canada 15  5
Examination of published data will show that some foods (e.g., Australia 20  3
rice and potatoes) have a wide variation in their GI values Canada 14  2
(Table 4). This is likely due to inherent botanical differences Sweet corn New Zealand 37  12
Australia 48
from country to country, compositional and processing differ-
China 55  2
ences, and possibly methodological differences in deriving GI
The United States 60
values (see Table 2). The majority of available GI values were South Africa 62  5
derived from commercially processed foods, and changes by
the food manufacturers in the ingredients or processing Source: Atkinson FS, Foster-Powell K, and Brand-Miller JC (2008) International tables
methods used will influence the GI. Manufactured foods with of glycemic index and glycemic load values: 2008. Diabetes Care, 31, 22812283.

Table 3 Method for calculation of the glycemic index of a mixed meal

Mathematical formula Practical example

Food GI CHO (g) Meal GI Food GI CHO (g) Meal GI

a GIa ga [GIa  ga]/G Roast beef 0.0 0.0

b GIb gb [GIb  gb]/G White bread 71 50.0 44.8
c GIc gc [GIc  gc]/G Peas and carrots 74 5.7 5.3
d GId gd [GId  gd]/G Margarine 0.0 0.0
e GIe ge [GIe  ge]/G Apple juice 38 23.6 11.3
Sum G Meal GI Sum 79.3 61.4

GI values are based on the glucose scale, using GI of glucose 100. CHO, AC.
Source: Wolever TMS (2013) Is glycaemic index (GI) a valid measure of carbohydrate quality? European Journal of Clinical Nutrition, 67, 522531.
246 CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits
and vegetables undergo ripening during which starch content
decreases and sugars increase; this will produce higher GI. Mois-
ture influences retrogradation and formation of resistant starch,
whereas cold storage after cooking can promote recrystallization
of the starch chains, which delays amylase activity. In this
regard, potato that was precooked, refrigerated overnight, and
reheated before consumption was shown to have a significantly
lower GI than potato that was cooked and eaten immediately.
Processing of grains and cereals disrupts the outer germ
layer, cell wall integrity, and starch granules, which in turn
influences starch particle size and exposes starch to enzymatic
digestion, thereby affecting GI. In the case of finely milled
particles, starch is more available to amylase compared to larger
or intact grains. Similarly, a given food might be processed
differently to produce different products (e.g., whole fruit,
whole fruit puree, and fruit juice). During the varied processing
methods needed to produce these items, there will likely be
changes in starch structures, particle size and relative abun-
dance of starch fractions; these will result in different GI values.
The influence of food fiber content on GI is not well
defined with many studies showing some or no effect. This
relationship is complicated by observations that soluble and
insoluble fiber appear to have varying effects on GI. In ran-
domized controlled trial (RCT), treatment with a low-GI diet
for 6 months resulted in lower HbA1c levels (a valid marker of
glycemic control) compared with a high-cereal fiber diet.
Examination of dietary interventions in persons with T2DM
shows that high-fiber diets improve glycemic control. How-
ever, it was apparent that those incorporating purified viscous
fibers more frequently lowered fasting blood glucose and total
cholesterol than interventions with purified nonviscous fibers.
These effects have been directly attributed to the viscosity of
purified fibers and the resulting effects on digestion and
absorption in the small intestine.
GI, GL, and Health Outcomes
Epidemiological Studies
Examination of epidemiological cohort studies in human
populations consistently shows an inverse association between
dietary whole grains and incident cardiovascular disease
(CVD). Current guidelines recommend that diets rich in min-
imally processed whole grains may lower risk for T2DM. How-
ever, the relationship between GI, GL, and T2DM seems to
differ by geographic location. In the United States, a recent
meta-analysis of three large studies confirmed previous find-
ings that dietary GI and GL are associated with increased risk of
T2DM. Further, the combination of low-GI or low-GL diets
with high cereal fiber intakes was associated with reduced
relative risk of T2DM. In fact, persons who consumed diets
with high GI or high GL, along with a low cereal fiber intake,
had 40% higher risk of T2DM than persons who consumed
low-GI or low-GL diets and had a high cereal fiber intake. This
analysis also showed that obese persons who consumed high-
GI or high-GL diets had a >10-fold higher risk of T2DM.
Further, a meta-analysis of ten studies on GI and T2DM and
14 on GL and T2DM, conducted in the United States (n 6),
Europe (n 4), and 1 each in the United Kingdom, China, and
Japan, showed that persons with the highest dietary GI and GL
had significantly higher relative risk of T2DM: 1.19 (CI
1.141.24) and 1.13 (CI 1.081.17), respectively. In contrast,
a cohort study in eight European countries found no relation-
ship between GI, GL, and T2DM. This difference may be attrib-
uted to the dietary assessment tools and methods used to assess
dietary intakes but may also reflect the limitations in using the
international table of GI and GL values. As discussed earlier,
inherent botanical differences and growing conditions of crops
and differences in formulation and composition of manufac-
tured goods may limit the generalizability of GI values derived
in one geographic region and used in another.
Randomized Controlled Trials
There are many limitations in examining dietdisease associa-
tion through epidemiological studies; this is overcome by con-
ducting randomized clinical trials, which is widely accepted as
an unbiased assessment of the effectiveness of an intervention.
Numerous RCTs have been conducted to examine the efficacy
of low-GI and low-GL diets in reducing risk factors for T2DM
and CVD and their effects on satiety and weight loss. Over the
past 5 years, several systematic reviews have been conducted in
order to assess the overall effectiveness of interventions with
low-GI and low-GL diets in persons with diabetes. These meta-
analyses showed that interventions of at least 4 weeks in dura-
tion consistently resulted in improved glycemic control, with
additional benefit seen when intake of unavailable carbohy-
drates is high. Similar reviews of existing RCTs have shown that
insulin sensitivity is improved by consumption of lower-GI,
lower-GL diets containing higher unavailable carbohydrates.
In terms of CVD risk factors, after adjusting for confound-
ing factors such as dietary fat intake, low GI was shown to
reduce fasting triacylglycerol, which is one of the two major
lipids found in blood. Conversely, high dietary GI is positively
associated with triglycerides and negatively associated with
HDLcholesterol. When legumes are included in low-GI die-
tary interventions, clinically significant reductions in HbA1c
are achieved, along with reductions in blood lipids, blood
pressure, and body weight. There is inconsistency in the rela-
tionship between low-GI or low-GL dietary interventions and
weight loss, although most studies show a nonsignificant trend
that favored these diets. When unavailable carbohydrate is
taken into account interventions with low GI or low GL, high
legume diets are significantly associated with reductions in
body weight and are probably linked to an associated reduc-
tion in energy intake. In fact, preliminary evidence suggests
that low-GI or low-GL diets may have superior satiety effects,
which may influence dietary energy intake.
Summary
GI can be used to classify the blood glucose raising potential of
carbohydrate-rich foods, but its derivation can be influenced
by many food factors. As such, several precautions must be
taken in order to obtain valid GI values. This could be achieved
within a regulatory framework where there is scrutiny over the
generation of GI values. Widespread utilization of GI in public
health promotion message has been limited due to concern
that increasing consumption of low-GI or low-GL diets could
 CARBOHYDRATES | Glycemic Index, Glycemic Load, and Their Health Benefits
lead to increased consumption of total and saturated fats,
thereby increasing CVD risk. However, recent evidence indi-
cates that lower AC intake following reductions in GI was not
accompanied with increased fat intake. Rather, they are associ-
ated with improved satiety and lower dietary energy intakes.
Exercises for Revision
Define glycemic index (GI) and explain the rationale for its
utilization.
Define glycemic load (GL) and explain the rationale for its
utilization.
Outline the method and precautions needed to derive the
GI value for a food.
Explain the derivation of the GI of a mixed meal.
Outline the food factors that may influence the GI value of
a food.
Discuss the precautions required to achieve accurate and
valid GI values.
Compare the findings of epidemiological and interven-
tional studies that have focused on the health benefits of
low-GI diets.
Exercises for Readers to Explore the Topic Further
What additional steps are required to improve the method
for GI so that it is universally accepted as being valid and
reproducible?
What is required to allow the widespread inclusion of GI
values on food labels?
How can GI values be better used for public health promo-
tion of healthy eating?
Is there a need for further human trials of low-GI foods? If
yes, what outcomes would be relevant?
What is the evidence that low-GI diets can influence brain
function, cognition, and satiety?
Acknowledgments
This article is supported by Agriculture and Agri-Food Canada
RBPI #1503, RBPI #2834/Agri-Innovation Program #068 with
Pulse Canada, and #T4.H29.V1 cluster program with Pulse
Canada.
Prof. Thomas Wolever kindly provided Table 3 and Figure 2.
Emily Padhi assisted with preparation of the other figures and
tables.
247
See also: Carbohydrates: Carbohydrate Metabolism; Cereals:
Chemistry and Physicochemistry of Non-starchy Polysaccharides;
Grains other than Cereals: Non-starch Polysaccharides; Resistant
Starch and Health; Food Grains and the Consumer: Grains and
Health; Grains and Health: Misinformation and Misconceptions; Food
Grains and Well-being: Functional Foods: Dietary Fibers,
Prebiotics, Probiotics, and Synbiotics.
Further Reading
Atkinson FS, Foster-Powell K, and Brand-Miller JC (2008) International tables of
glycemic index and glycemic load values: 2008. Diabetes Care 31: 22812283.
Bhupathiraju SN, Tobias DK, Malik VS, et al. (2014) Glycemic index, glycemic load, and
risk of type 2 diabetes: Results from 3 large US cohorts and an updated meta-
analysis. The American Journal of Clinical Nutrition 100: 218232. http://dx.doi.
org/10.39345/ajcn-079533.
Fabricatore AN, Ebbeling CB, Wadden TA, and Ludwig DS (2011) Continuous glucose
monitoring to assess the ecologic validity of dietary glycemic index and glycemic
load. The American Journal of Clinical Nutrition 94(6): 15191524.
Food and Agriculture Organization of the United Nations (1998) Carbohydrates in
Human Nutrition. Rome: Food and Agriculture Organization of the United Nations
Food and Nutrition, Paper 66.
International Organization for Standardization (2010) Food products Determination
of the glycaemic index (GI) and recommendation for food classification.
International Organization for Standardization, September 23, 2010 (ISO
26642:2010). Available from: http://www.iso.org/iso/catalogue_detail.htm?
csnumber43633 (cited 18 August 2014).
Jenkins DJA, Wolever TMS, Taylor RH, et al. (1981) Glycemic index of foods:
A physiological basis for carbohydrate exchange. American Journal of Clinical
Nutrition 34: 362366.
Lan-Pidhainy X and Wolever TMS (2011) Are the glycemic and insulinemic index
values of carbohydrate foods similar in healthy control, hyperinsulinemic and type 2
diabetic patients? European Journal of Clinical Nutrition 65: 727734.
Li SS, Kendall CW, Souza RJ, et al. (2014) Dietary pulses, satiety, and food intake: A
systematic review and meta-analysis of acute feeding trials. Obesity (Silver Spring)
22(8): 17731780. http://dx.doi.org/10.1002/oby.20782.
Livesey G, Taylor R, Hulshof T, and Howlett J (2008) Glycemic response and health
A systematic review and meta-analysis: Relations between dietary glycemic
properties and health outcomes. The American Journal of Clinical Nutrition 87(1):
258S268S.
Mirrahimi A, Chiavaroli L, Srichaikul K, et al. (2014) The role of glycemic index and
glycemic load in cardiovascular disease and its risk factors: A review of the recent
literature. Current Atherosclerosis Reports 16(1): 110.
Salmeron J, Manson JE, Stampfer MJ, Colditz GA, Wing AL, and Willett WC (1997)
Dietary fiber, glycemic load, and risk of non-insulin-dependent diabetes mellitus in
women. JAMA 277: 472477.
Sluijs I, Beulens JW, van der Schouw YT, et al. (2013) Dietary glycemic index,
glycemic load, and digestible carbohydrate intake are not associated with risk
of type 2 diabetes in eight European countries. Journal of Nutrition 143(1):
9399.
Venn BJ and Green TJ (2007) Glycemic index and glycemic load: Measurement issues
and their effect on dietdisease relationships. European Journal of Clinical Nutrition
61(Suppl. 1): S122S131.
Wolever TMS (2013) Is glycaemic index (GI) a valid measure of carbohydrate quality?
European Journal of Clinical Nutrition 67: 522531.
Wolever TMS, Brand-Miller JC, Abernethy J, et al. (2008) Measuring the glycemic index
of foods: Interlaboratory study. The American Journal of Clinical Nutrition 87(1):
S247S257.
 Fats
Contents
Lipid Chemistry
Healthy Fats and Oils
Lipid Chemistry
L Day, AgResearch Ltd., Palmerston North, New Zealand
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The major lipid class in cereal grains and oilseeds is
triacylglycerols.
The minor complex lipids are phospholipids, glycoglycer-
olipids and sterols.
The composition of the lipids (or oil) in oilseeds deter-
mines its food or non-food applications.
Extraction and quantification of different lipid classes are
achieved by solvent selections and various chromatograph-
ic methods.
Supercritical fluid carbon dioxide with or without cosolvent
is now applied for lipid analyses and for industrial extrac-
tion of oils.
Mass spectrometry is increasingly being used for the deter-
mination of lipids.
Lipids play an important technological role in cereal pro-
duction, storage, processing and quality of cereal-based
foods.
Plant lipids contain more essential polyunsaturated fatty
acids than animal fats, and their inclusion in food is posi-
tive from a nutritional point of view.
Learning Objective
To understand lipid classes in grains and oilseeds, their
analysis, their technological properties in processing, and
nutritional values of plant lipids.
Introduction
After protein and carbohydrate, lipids (commonly known as
fats or oils) are the third most important macronutrient in
human and animal diets. Although lipids are relatively minor
constituents in food grains, they play an important technolog-
ical role in the quality of grains upon storage: and in most
aspects of processing, including milling, brewing, baking and
248 Encyclopedia of Food Grains, Second Edition
extrusion. On the other hand, oilseeds are primarily grown for
their oil content. Thus, the composition of the lipids (or oil) in
various oilseeds is an essential consideration in determining
the use of the oil. Lipids are also essential for the nutrition and
consumer acceptance of finished food products as they are
carriers of fat-soluble flavor compounds and vitamins. The
terminology of lipid chemistry is largely dependent on the
extraction conditions, including the solvents used and temper-
ature, and quantification methodologies.
Classification
Lipids can be classified based on their physical properties at
room temperature (oils are liquid and fats are solid) or their
polarity (neutral lipids include fatty acids (FAs), glycerides, and
sterols and polar lipids include glycerophospholipids (phos-
pholipids) and glyceroglycolipids (glycolipids)). The most com-
mon classification of lipids is based on their structure simple
or complex. Simple lipids, composed of FAs and alcohol,
include acylglycerols, ether acylglycerols, sterols and their esters
that can be hydrolyzed to two different components usually an
alcohol and an acid. Complex lipids include phospholipids,
glycolipids and sphingolipids, which yield three or more differ-
ent compounds on hydrolysis.
The FAs (carboxylic acids) are the main building blocks in
lipid structures. Only a small portion of the total lipid fraction
consists of free carboxylic acids, or free fatty acids (FFAs). Most
of the carboxylic acids in the lipid fraction are found as esters of
glycerol, that is, as triacylglycerols (TAGs), commonly known
as triglycerides (Figure 1).
The most common TAGs are those with long-chain carbox-
ylic acids (C14C22). The positions of carboxylic acids in TAGs
are also important to their properties and utilization. The
saturated forms of FAs are the most stable against oxidation.
They also have high melting temperatures in comparison with
unsaturated FAs with the same number of carbon atoms. In
plant TAGs, unsaturated FAs are predominantly located in the
2 position. Monoacylglycerols (MAGs) contain only one long-
http://dx.doi.org/10.1016/B978-0-12-394437-5.00099-1
 FATS | Lipid Chemistry 249

Glycerol
Ester Compositions and Distribution of Lipids in Cereal
Carbon
Linkage Grains and Oilseeds
Position
H O H O
Cereal Grains
1 H C OH + H O C R H C O C R
O O Lipids are minor components of the cereal grains. Wheat,
2 H C OH + H O C R H C O C R
barley, brown rice, rye and sorghum have lower lipid contents
O O than other cereal crops such as oats, pearl millet, and maize. All
cereal grain lipids are richer in nonpolar lipids than in polar
3 H C OH + H O C R H C O C R
lipids. Wheat contains the highest level of glycolipids among
H H all cereal grains, followed by triticale, rye, and barley; millet,
Glycerol Fatty acid Triglyceride maize, and sorghum lipids contain the lowest glycolipid con-
Figure 1 Triacylglycerol synthesis: R, R0 , R00 are usually long-chain
tent. In general, phospholipids are more abundant in wheat,
alkyl groups and R, R0 , R00 are typically all different and may also contain triticale and rye lipids and slightly lower in the lipids of barley,
one or more carboncarbon double bonds. maize, oat, sorghum and rice. Approximate lipid contents and
the proportions of nonpolar lipids and polar lipids (separated
into glycolipids and phospholipids) in each cereal are listed in
Table 1 General classification of lipids and their commonly used Table 2.
abbreviations TAGs are the major nonpolar lipids in cereal grains, repre-
senting from
50% of the total nonpolar lipids in wheat and
Lipid class Abbreviation
barley to as high as 90% in maize. They are deposited in
Simple lipids: compounds with two types of structural spherosomes (oil droplets) bound by a monolayer membrane
moiety and are usually stored by plants in this form. The remainder of
Glyceryl esters: esters of glycerol and fatty acids (see the nonpolar lipids are mainly di- and monoglycerides, FFAs,
Figure 1) and sterol esters. The major phospholipids in cereal grains are
Triacylglycerols: 3  fatty acids TAGs phosphatidylcholine (PC), phosphatidylethanolamine (PE),
Diacylglycerols: 2  fatty acids at any 2 positions DAGs and phosphatidylinositol (PI). Monoacyl PLs, or lysophospho-
Monoacylglycerols: 1  fatty acid at any position MAGs lipids (prefix L added to LP abbreviation, Table 1), are usually
Sterol esters: esters of sterol and fatty acids SEs regarded as degradation products of PL. The GLs are quantita-
Waxes: esters of long-chain alcohols and fatty acids
tively the major components of the glycolipids in whole cereal
Complex lipids: compounds with more than two types
of structural moiety
grains and starchy endosperm. The principal sugar in the GLs is
Phospholipids: glycerol esters of fatty acids, galactose, and glucose is either a minor component or absent.
phosphoric acid, and other groups containing The main galactosylglycerides are monogalactosyldiglyceride
nitrogen (MGDG) and digalactosyldiglyceride (DGDG). Small amounts
Phosphatidic acid: diacylglycerol esterified to of galactosylmonoglycerides (monogalactosylmonoglycerides
phosphoric acid (MGMG)) and digalactosylmonoglycerides (DGMG) are also
Phosphatidylcholine: phosphatidic acid linked to PC found in the endosperm of some mature cereal grains. Polar
choline lipids are found in all membranes including the amyloplast
Phosphatidylethanolamine PE membrane. Apart from these major acyl lipids, the other lipids
Phosphatidylserine PS
in cereals include sterols and lipid-associated compounds such
Phosphatidylinositol PI
Lysophosphatidylcholine, LPC, LPE,
as carotenoids and tocopherols. Although carotenoids are very
lysophosphatidylethanolamine, LPS, LPI minor constituents in cereal grains, color contributed by carot-
lysophosphatidylserine, lysophosphatidylinositol enoids is an important factor in the use of cereal grains in food
Glycoglycerolipids: 1,2-diacylglycerol joined by a production, particularly in the use of durum wheat for pasta
glycosidic linkage through position sn-3 to a making.
carbohydrate moiety The lipid content in cereals is influenced by the water
Monogalactosyldiglyceride MGDG content, the stage of maturity, and the variety of the grain
Digalactosyldiglyceride DGDG crop. Lipids are also unevenly distributed in various parts of
Monogalactosylmonoglyceride MGMG cereal grains. For example, in the wheat grain, the germ com-
Digalactosylmonoglyceride DGMG
prises only
4% of the total grain, by weight, but it has the
highest lipid content and the highest proportion of polar lipids
(Table 3), whereas the endosperm, the major fraction of wheat
grain, has significantly lower lipid content than the other
chain FA (saturated or unsaturated) at either the 1, or 2, or 3 fractions. Lipids in flour (except whole meal) can be subdi-
position. Similarly, diacylglycerols (DAGs) have two FA chains vided into nonstarch and starch lipids (Figure 2). The non-
at any of the two positions. However, the phosphoric acid starch lipids are further divided into free lipids and bound
residue of phosphoglycerides (PLs) and the first glycosidic lipids based on the extractability of lipids by solvents. The
residue of glycosylglycerides (GLs) are always in the 3 position. lipids that are easily extractable from flour or freeze-dried
The general classification of the principal lipids found in cereal dough, with a nonpolar solvent such as hexane, ether, or
grains and oilseeds, brief descriptions of their basic structures, petroleum ether, are called free lipids. Bound lipids refer to
and their common abbreviations are listed in Table 1. the nonstarch lipids that are only extractable with a very polar
250 FATS | Lipid Chemistry

Table 2 Approximate lipid contents and the distribution of lipid classes of the total lipids in cereal grains

Lipid class (% w/w of TL)

Cereal grains Total lipids (% w/w in whole grain) NL GL PL

Barley 2.54.7 6578 726 926

Maize/corn 5.16.0 8896 3 5
Millet 1.711.0 7594 115 114
Oats 4.510.3 6680 610 1226
Rice (brown) 0.93.1 7887 412 810
Rye 3.5 6371 1012 1825
Sorghum 3.66.0 7786 26 1117
Triticale 2.64.6 5367 1018 1729
Wheat 2.13.8 6072 1222 1426

TL, total lipids; NL, nonpolar lipid; GL, glycolipid; PL, phospholipid.

Table 3 Distribution and compositions of lipids in wheat grain fractions

Lipid class (% w/w of TL)

Grain fraction Grain composition (%) Lipid distribution (%) Total lipids in each fraction (% w/w) NL GL PL

Whole grain 100 100 2.13.8 6072 1222 1426

Germ 4 3035 2535 7985 03.5 1417
Bran 4 25 24 7280 610 1318
Endosperm 92 6070 1.52.5
Nonstarch lipid 3346 3038 2434
Starch lipid 46 15 9095

TL, total lipids; NL, nonpolar lipid; GL, glycolipid; PL, phospholipid.

Wheat flour lipids PL, whereas starch lipids are predominantly PL and almost
exclusively lyso-PL in which LPC is the main lipid component.
The distribution of lipid classes in other cereal grains is
Nonstarch lipids Starch lipids qualitatively similar to those in wheat. The germ is the richest
(1.4%) (1.0%) source of lipids among all cereal grain fractions; more than
one-third of the total lipids is in the germ. In particular, maize
has a larger proportion of germ in the grain, 1125%, by
Free lipids Bound lipids
weight, with an exceptionally high lipid content, 3947% by
(0.8%) (0.6%)
weight, containing mostly TAGs, with a small amount of PL
and traces of GL. Maize germ is, therefore, more suited for
Nonpolar lipids Polar lipids
(<0.1%) (0.9%) edible oil production than any other cereal, after it has been
Nonpolar lipids Polar lipids
(0.6%) (0.2%) separated from the starch endosperm.
The FA composition of cereal lipids is generally similar for
wheat, barley, triticale, maize, and sorghum (Table 4). All
TAG, DAG, MAG, GL, PL LPC cereal grain lipids are rich in unsaturated FAs. Palmitic acid
FFA, SE (16:0) is a major saturated FA, and linoleic acid (18:2) is a
major unsaturated FA for all cereals expect for brown rice. Rye
Figure 2 The subdivision of wheat flour lipids. TAGs, triacylglycerols;
DAGs, diacylglycerols; MAGs, monoacylglycerols; SEs, sterol esters; lipids contain slightly higher levels of linolenic acid (18:3)
GLs, glycolipids; PLs, phospholipids; LPC, lysophosphatidylcholine. than those of other cereals. Oat lipids are similar to those of
brown rice; both are rich in oleic acid (18:1).

solvent such as chloroform or water-saturated n-butanol from

the flour with free lipids already removed. When free and
bound lipids are extracted from flour and freeze-dried dough,
the free lipid fraction in dough is only one-third of the amount
of flour-free lipids. This is because two-thirds of the flour lipids
become bound, mainly to gluten during dough mixing. The
nonstarch lipids, which consist of all the endosperm lipids,
excluding those inside starch granules, contain more GL than
Oilseeds
Unlike the cereal grains, lipids (oils) are the main constituent
of the oilseed crops. Oilseeds such as canola/rapeseed, saf-
flower and sunflower are grown for their oil. On the other
hand, oil is produced as a by-product from other crops such
as soybeans, which are grown primarily for their protein-rich
content, and cottonseed grown for its fiber. Peanut, a
 FATS | Lipid Chemistry 251

Table 4 Common FA compositions of total lipids in cereal grains

Fatty acid (wt%)

Lipid source Palmitic (16:0) Stearic (18:0) Oleic (18:1) Linoleic (18:2) Linolenic (18:3)

Wheat 1724 12 821 5560 35

Barley 2124 12 914 5659 47
Oats 726 14 2248 3152 14
Rye 1219 12 1216 5765 312
Maize/corn 1137 15 1124 4663 16
Brown rice 1426 13 3652 2539 14
Sorghum 1017 12 2127 4258 15
Triticale 1620 12 814 5760 46

Table 5 Oil contents in oilseeds
Oilseeds Oil in dry seed (% w/w)
Canola 4050
Rapeseed/mustard 3050
Cottonseed 1525
Linseed 3545
Peanut 3656
Safflower 1247
Soybean 1522
Sunflower 2550
leguminous oilseed, is used not only for edible oil production
but also for direct consumption in various human foods, being
a good source of protein and oil. Another oilseed crop, linseed,
produces oil that is not extensively used for food, but it is
nevertheless an important industrial oil.
Lipid contents vary among the oilseed crops, shown as oil
contents in Table 5, ranging from about 15% to 25% in soy-
bean and cottonseed, to as high as 50% in sunflower and
canola/rapeseed, to 56% in peanut. It must be emphasized
that both oil content and component FA proportions in an
oilseed are subject to quite a wide variation. Such variations
arise from genetic manipulation of seed varieties, climatic fac-
tors and cultivation practices.
Lipids in oilseeds are predominantly TAG,
90% or more
of total lipids, and phospholipids being the main polar lipids.
Other minor lipid constituents are FFAs, sterols, hydrocarbons,
etc. The FA composition of lipids is the main influence in terms
of the quality and the uses of the oil. Cottonseed oil has high
levels of saturated FAs among the oilseeds, 1825% palmitic
acid, and a small amount (13%) of myristic acid (Table 6).
The main unsaturated FAs are linoleic acid (3252%) followed
by oleic acid (1032%). Cottonseed oil has a significant stor-
age stability not only due to its lesser amount of unsaturated
FAs content but also due to the presence of enough tocopherols
to reduce lipid peroxidation.
Among the common oilseeds, safflower, sunflower, and
peanut oils contain high levels of unsaturated FAs. Sunflower
oil is characterized by its high concentration of linoleic acid
(6070%) followed by oleic acid (1535%). The high propor-
tion of polyunsaturated FAs makes sunflower oil a popular
source of essential FAs in the diet. Sunflower, safflower and
soybean oils are the oils containing the highest levels of
linoleic acid (C18:2). The FA composition of safflower oil is
similar to that of sunflower. The oil of commercial safflower
cultivars contains 5581% linoleic acid and 742% oleic acid
as major FAs, followed by stearic acid (110%) and palmitic
acid (210%) as minor FAs. In peanut oil, oleic and linoleic
acids constitute 3371% and 1246% of the total FAs, respec-
tively. A high proportion of unsaturated FAs in an edible oil,
however, does decrease its storage stability and quality, due to
oxidation reactions associated with the double bonds in unsat-
urated FAs. In addition, more and more nutritional studies are
showing that linoleic acid is not necessarily good for your
health. A new type of sunflower that has about 6571% oleic
acid has been produced to replace the traditional sunflower.
Unlike other oilseeds, FA composition in rapeseed oil is
unusual in that it contains substantial amounts of long-chain
FAs, in particular a significant amount of erucic acid (22:1) as

2050% of total lipids (Table 6). Major changes in the FA
composition of rapeseed oils have arisen as a result of selective
breeding for low levels of erucic acid. This is because of some
indications that erucic acid had antinutritional properties.
Consequently, varieties of rapeseed (called canola) have
been developed in which the level of erucic acid is virtually
zero. Canola varieties are also characterized by markedly
increased levels of oleic acid with smaller increases also in
linoleic and linolenic acid content compared with those of
the former rapeseed varieties due to the substantial reduction
of erucic acid content (Table 6). Canola is the oil with the
lowest content of saturated FAs. Linolenic acid content has also
been reduced in some canola varieties in favor of linoleic acid,
through selective breeding. Low-linolenic acid content canola
varieties could have less than 2% linolenic acid and resemble
olive oil in terms of their monounsaturated FA contents more
than the high linolenic acid content oil variety (conventional
canola varieties).
Linseed oil is characterized by high levels of linolenic acid
(5260%), which undergoes autoxidation to give thin, cross-
linked, and tough films, making it unsuitable for edible pur-
poses. It is, however, an important drying oil used to produce
oil-based paints.
Extraction and Quantification of Lipids
Accurate and precise analyses of lipids are important, not only
for the determination of their nutritive values for human
252 FATS | Lipid Chemistry
Table 6 FA compositions of total lipids in oilseeds
Fatty acid (% w/w)
Lipid source Palmitic (16:0) Stearic (18:0) Oleic (18:1)
Canola 35 11.5 5760
Rapeseeds 13 0.71 1214
Cottonseed 1825 12 1738
Linseed 57 34 1820
Peanut 814 26 3756
Safflower 210 110 742
Soybean 612 25 2034
Sunflower 57 26 1535
health benefits, but also for the understanding of the structural
characteristics and functional properties of lipids, which may
allow the development of tailor-made products that are
designed for a particular function or application. Analyses of
lipids involve several steps which usually include preparation
of the sample, solvent extraction of the lipids, removal of the
solvent and separation of the lipids into classes for qualitative
or quantitative analyses. Sample preparation mainly involves
particle-size reduction of plant tissues or seed materials using
methods such as grinding. The purpose of this step is to
increase surface area and facilitate subsequent lipid extraction.
In some cases, acid hydrolysis may be required to release lipids
which are covalently bound to proteins and carbohydrates.
Solvent Extraction
The choice of organic solvents for lipid extraction largely
depends on both the chemical nature of the sample and the
type of lipids to be extracted, that is, total lipids, starch lipids,
and polar or nonpolar lipids. Neutral lipids are hydrophobi-
cally bound and can be extracted readily by nonpolar solvents,
whereas polar lipids, such as glycolipids and phospholipids,
which are predominantly bound by electrostatic forces and
hydrogen bonding, require polar solvents capable of breaking
such bonds. Hexane (petroleum ether) is the preferred solvent
for extracting nonpolar, neutral lipids such as triglycerides and
is the main solvent used for oilseed crushing/extraction for the
industrial production of crude oil from oilseeds because of its
low cost. However, cereal grains and milled flour contain
relatively large amounts of glycolipids and phospholipids,
and large amounts of cereal lipids (about half of the total
lipids) are bound to polypeptides or polysaccharides. There-
fore, more polar solvents such as chloroform, alcohol, or
chloroformmethanol (2:1) mixture are needed for extraction.
Hexane combined with an acid hydrolysis will give a complete
extraction of the lipids from cereals.
Removal of the solvent from lipid extracts is normally con-
ducted under vacuum in a rotary evaporator at or near room
temperature. Care must be taken to minimize lipid oxidation
during sample preparation, solvent extraction and storage of
lipids. Cereal lipids characteristically contain large proportions
of polyunsaturated FAs together with carotenoids and tocols,
all of which are highly susceptible to autoxidation. The lipids
are only slowly oxidized while the grain tissues are intact, but
once the tissues have been disrupted, oxidation must be
Linoleic (18:2) Linolenic (18:3) Others
1020 1012
1122 810 2050% erucic (22:1)
4555 13% myristic (14:0)
1517 5260
2540
5581
4656 512
6073
minimized by keeping lipids in solvents such as chloroform
and under a nitrogen atmosphere as well as by working at low
temperatures if possible.
Supercritical Fluid Extraction
Supercritical fluid extraction (SFE), which utilizes the solvation
power of the gases when compressed above critical point fluids
at pressure and temperature, has been applied for lipid ana-
lyses and for industrial extraction of oils. Carbon dioxide
(CO2) is the most widely used supercritical fluid. The prepara-
tion of the sample before extraction is extremely important. For
solid materials, for example, grains, seeds, flours, or meals, the
material needs to be ground to produce smaller particles in
order to increase surface area and promote more efficient
extractions. Supercritical CO2 is generally considered a more
selective solvent than hexane. The fact that supercritical CO2 is
a good solvent for nonpolar and moderately polar compounds
makes it an excellent choice for extracting lipids without
extracting other more polar nonfat components. In this way,
it is possible to extract nonpolar TAG without the coextraction
of PL. However, polar lipids such as PL can also be extracted by
the supercritical CO2 method with the use of solvent modifiers
such as ethanol, which increases the polarity of the supercrit-
ical CO2. The SFE method uses no hazardous solvents, so it is
well suited for the process-control environment.
The SFE-based test methods have been approved as an
official method by AOCS for the determination of process-
scale extractable oil or determination of total oil. The method
has two parts involving extraction with supercritical CO2 alone
or plus 15% ethanol. Although SFE has been used for wheat
lipid extraction, there is no official SEF-based method for
extraction of lipids from wheat and other similar cereal grains.
Lipid extracts are complex mixtures of individual classes of
compounds and require further separation. Many chromato-
graphic and spectroscopic techniques are in use to separate and
quantify individual lipid components. The most commonly
used techniques are thin-layer chromatography (TLC), high-
performance liquid chromatography (HPLC) and gas chroma-
tography (GC).
Thin-Layer Chromatography
TLC provides the separation of a wide variety of compounds
with different polarities on a single plate and has long been

Triglycerides
Free fatty acids
Glycolipids
Phospholipids
1 2 3 out either with a solvent mixture of constant composition or
Figure 3 Separation of wheat flour lipids by TLC on a silica gel G plate
(20  20 cm) developed in chloroform/methanol/water (90:20:2 v/v/v).
Wheat flour was collected from different mill streams (13: earlylate),
where more TAGs were detected in the later mill streams as progressively
more bran materials were being incorporated into flour.
used for the separation of cereal lipids. Silica is still the station-
ary phase material preferred. Small amounts of lipid material
are spotted on a plate. The plate is then developed in a lid-
covered glass chamber containing a mixed solvent system,
which behaves like a gradient elution during travel on a
plate. The separation of lipids is based on relative affinities of
the components to the TLC adsorbent. When the solvent front
is close to the top of the plate, the plate is removed from the
chamber and residual solvent is evaporated under an inert
atmosphere. The separated lipid spots can be visualized as
brown spots (temporarily) by exposing to iodine vapor or
under UV light after spraying with fluorescent agents or as
dark spots (permanent) by spraying with sulfuric acid
(1050% in ethanol v/v) and subsequently charring above
160  C. The advantage of using a temporary nondestructive
detection is that the lipid spots may be recovered from the
silica gel for preparative or analytic proposes. Lipids, after
separation on a silica plate, may also be quantitated by densi-
tometric or fluorimetric scanning.
Cereal lipids can be separated into various classes by TLC.
As an example, Figure 3 shows the separation of wheat flour
lipids extracted by chloroform from different flour mill
streams, where more TAGs were detected in the later mill
streams (lane 3) as more bran materials were being incorpo-
rated into flour. Straightforward unidimensional TLC is ade-
quate for the separation of nonpolar lipids and some polar
FATS | Lipid Chemistry 253
lipids. In most cases, glycolipids overlap phospholipids in TLC
systems that separate phospholipids. More complex GLs or PLs
are preferentially separated by two-dimensional TLC, using a
neutral/basic solvent such as in the first dimension and an
acidic solvent in the second dimension.
TLC offers a much higher sample throughput due to the
possibility of performing separations simultaneously, and it
can handle not-so-clean samples since the separation medium
is used only once. TLC is a cost-effective, simple, and easy
chromatographic technique.
High-Performance Liquid Chromatography
In principle, separations by HPLC are carried out by essentially
the same separation principle as TLC. Using HPLC, it is easier
to generate large numbers of theoretical plates (a measure of
resolution efficiency); HPLC is simpler to automate than TLC.
However, HPLC is much more expensive than TLC in terms of
both equipment and running costs. HPLC separation occurs in
a stainless steel column packed with a very uniform, finely
divided, microspherical adsorbent material of controlled
porosity and degree of hydration. A high-pressure pump
ensures adequate and constant flow of solvent, the mobile
phase, through the column, and a flow-through detector con-
tinuously monitors the column eluate. Adsorption chromatog-
raphy with columns of silica gel is commonly used for the
separation of lipids. Elution of the column may be carried
by gradient elution, in which the solvent composition is varied
linearly or in a stepwise fashion with both binary and ternary
solvent systems. The choice of the elution systems is sometimes
restricted by the use of the detection system. For example, a
solvent mixture of constant composition with low UV absor-
bance may be essential for a UV detector. Although gradient
solvent systems can be used with an evaporative light-
scattering detector (ELSD), the choice of solvents is con-
strained by the need for sufficient volatility for evaporation in
the detector under conditions that do not cause evaporation of
the solute. Complicated programs (up to eight steps) are some-
times required to achieve the desired separations. However,
ELSD has presently become the detection method used most
often in the separation of lipids by HPLC. HPLC for analysis of
all simple lipid classes in one chromatographic run has not yet
been used widely, probably because TAGs (major components
in all cereal grains and oilseeds) dominate the system. HPLC
methods have been devised more for specific analysis of indi-
vidual components. The use of HPLC for the separation of
complex lipids has increased. Separation of glycolipids
MGDG and DGDG, along with several phospholipids from
wheat flour lipid extracts at analytic scale (Figure 4), as well
as preparative scale, has been achieved.
Gas Chromatography
GC is the most useful tool for the analysis of the FA composi-
tion in lipids. Derivatization of FAs is required to increase their
volatility. Fatty acid methyl esters (FAMEs) may be prepared by
different transmethylation techniques and then separated on
GC columns and detected by a flame ionization detector (FID).
The gas phase for GC is usually nitrogen, hydrogen, or helium.
254 FATS | Lipid Chemistry
DGDG
1200
1000
Response (mV)
MGDG
800
600
400
PG
200
PC
0 100
10 15 20 25 30 35 40
Time (min)
Figure 4 HPLC analysis of wheat flour polar lipids extracted by
chloroform. Separation was carried out on a PVA-Sil threaded guard
column (23  4.0 mm i.d.) with gradient elution at 0.5 ml min1 and
detected by a ELSD. Four steps of gradient were used for the separation
010 min, 100% eluent A (2,2,4-trimethylpentane); 1020 min, 90%
eluent A 10% eluent B (tert-butylmethylether); 2035 min, 100%
eluent B; and 3540 min, 100% eluent C (methanol/n-ethylmorpholine/
glacial acetic acid (500:4.2:1.5 v/v)). For abbreviation of lipids, see
Table 1.
Since the principal FAMEs in cereal and oilseed lipids are 16:0,
16:1, 18:0, 18:1, 18:2, and 18:3, heptadecanoic acid (17:0) can
be used as an internal standard. Identification and quantifica-
tion of individual FAs can be achieved by comparing and
calibrating with relatively cheap FAME standards. Figure 5
shows a typical GC profile of FA compositions in canola oil.
FA compositions in lipid classes can also be obtained by GC
analysis of lipids in each class separated or prepared by TLC or
other column chromatographic techniques.
Lastly, mass spectrometric detection is being used increas-
ingly for the detection and analysis of complex lipids. The
technique can be used with direct inlet injection or linked to
HPLC or GC systems where the mass spectrometer serves both
to detect lipids and to obtain information on their molecular
structure.
Functionality of Lipids in Bakery Products
Apart from being used directly as a source of human food and
animal feed, a large proportion of cereal grains are milled to
flour, which is then used for producing various food products.
Wheat has been the predominant cereal grain used for baked
goods, largely due to its ability to form dough when flour is
mixed with water. Rye ranks next in importance while other
cereals assume a role in specialized products and in the prep-
aration of composite flours.
The rheological properties of doughs made from wheat
flour are principally determined by the gluten protein, and
lipids (naturally occur in cereal grain, excluding added fat)
appear to make little contribution. When protein is removed
from flour, dough and bread-making properties are lost. In
contrast, good dough properties and bread-making capacity
are retained after removal of the nonstarch lipids. However,
900
18:1 oleic
800
700
Response (mV)
600
18:2 linoleic
500
400
18:3 linolenic
16:1 palmitic
18:0 stearic
300
200
0
20 25 30 35
Time (min)
Figure 5 GC analysis of a canola oil. FAs methyl esters separated using
a capillary GC column (SP2560, Supelco) and detected by an FID at
240  C.
flour lipids do affect the loaf volume and crumb texture of
bread. Fractions of polar and nonpolar lipids have opposite
effects on baking performance.
Incremental addition of the extracted lipid to a defatted
flour produces an unusual effect on the loaf volume and tex-
ture in a rapid baking test in which exogenous lipid is omitted.
Loaf volume decreases to a minimum at a lipid content inter-
mediate between those of the defatted flour and untreated
flour and thereafter increases. At high lipid contents (above
the inherent lipid content of the flour), the loaf volumelipid
curve tends to plateau. The changes in volume are paralleled by
changes in crumb grain, which, at first, also deteriorates and
then improves. Fractions of polar and nonpolar lipids affect
loaf volume and crumb grain in opposite ways. The polar lipid
fraction has generally beneficial effects, whereas the nonpolar
fraction has deleterious effects. Thus, if the ratio of polar
to nonpolar lipid is increased, the minimum of the loaf
volumelipid content curve is shifted to lower lipid contents.
If the percentage of polar to nonpolar lipid in a flour is varied
from 0 to 100 at a constant lipid level, test-bake loaf volume
increases approximately in a linear manner. The addition of
different lipids to test their effects on dough has shown that the
unsaturated FAs such as linoleic acid are the components that
mainly contribute to the deleterious effects of the nonpolar
fraction. Higher loaf volume and better crumb texture in bread
are favored by a high ratio of polar/nonpolar lipid and a high
content of flour lipid. However, the variation in loaf volume
that can be attributed to lipid is relatively small, and by far, the
greatest variation is imposed by the effect of gluten protein
quality.
The role of the polar lipid appears to act as a complex
mixture of components with surface activities in stabilizing or
destabilizing the gas bubble structure during expansion of the
loaf and thus the final texture. Upon heating (i.e., during
baking), the dispersed lipids provide an interfacial film at the
surface of gas cells, thus helping to modulate the growth,
expansion and coalescence of air cells. In addition to their
effects in influencing volume and texture of baked products,
lipids also play a role in staling mechanisms. Generally, the

presence of lipid reduces bread staling and enhances shelf life.
This is largely due to the interaction between lipids and starch-
es. When monoacyl lipid monomers are present to form the
lipidstarch complex, an insoluble amyloselipid film is likely
to be formed on the surface of the starch granules. Such a film
is capable of acting as a barrier against water transport involved
in the staling process.
The effects of lipids have also been studied for Arabic bread,
steamed bread, cakes and biscuits. Despite differences in pro-
cessing, the general features with respect to lipids have much in
common. It is the crumb texture that is influenced most, and,
like bread, differences in lipids have not been found to account
for major variations in quality. However, the contribution of
the lipid component to flour quality should not be ignored
when considering parameters to be used as a basis for selection
in plant breeding programs even though protein composition
appears to be the major factor that determines flour quality.
The quality factors in flour lipid, as presently known, are a
high nonstarch lipid content, a high ratio of polar to nonpolar
lipid, and a low free fatty acid content.
Nutritional Values
Lipids are the richest source of energy on a weight basis. They
are essential components of our diet and play vital physiolog-
ical and biochemical roles in human health and well-being.
Lipids also increase palatability, including taste, aroma, and
texture of food, because they are carriers of fat-soluble flavors.
Fats and oils are also good carriers of fat-soluble vitamins such
as vitamins A, D, E, and K. Plant lipids contain tocopherols and
tocotrienols, which are the most important sources of vitamin
E. Appropriate intake of lipids is essential for health mainte-
nance. However, a high consumption of fat, especially satu-
rated FAs, may be connected with several chronic diseases, such
as heart disease and obesity.
Of the lipid components of a normal diet, the most impor-
tant FAs are linoleic acid (n-6) and a-linolenic acid (n-3), the
two primary essential polyunsaturated fatty acids (PUFAs).
They play a role in stabilizing biological membranes by creat-
ing physical properties that are optimal for the transport of
substances across the membrane and for the biochemical reac-
tions occurring in the membrane. Through metabolism, they
are converted into a whole range of oxygenated compounds
which exert a range of profound physiological activities involv-
ing lowering plasma cholesterol, aggregating red blood cells,
and smoothing muscle performance, all attributes that are
required for good health. However, studies have shown that
too much linoleic acid (C18:2) can have adverse effects on
health, especially causing inflammatory conditions. Since the
human body is unable to synthesize them, they must be
obtained from dietary sources. Since plant lipids contain more
essential PUFAs than animal fats, their inclusion in food is
positive from a nutritional point of view. The important source
of the essential FAs is oilseeds. Although cereal grains contain
high amounts of linoleic acid, due to the generally low total
lipid content of the grain, they are unlikely to have much contri-
bution to the essential FA intake in the human diet, except when
the lipids are extracted from the lipid-rich germ and produced
as oil, such as maize, or rice germ oil or wheat bran oil.
FATS | Lipid Chemistry 255
Oils extracted from oilseeds, however, have a sufficient influence
in the total fat intake of the human diet since they are used
directly for frying foods, as cooking ingredients, and as spreads.
Recent developments in plant molecular biology now allow the
engineering of oilseeds with increased omega-3 long-chain
PUFAs through complex multigene biosynthetic pathways.
Such development can have a considerable potential to increase
the nutritional values of plant-based oils.
Exercises for Revision
What are the different classes of lipids in cereal grains and
oilseeds?
What are the most common fatty acids in cereal grains and
oilseeds?
What are the different solvents used for the extraction of
nonpolar or polar lipids?
What is the most common gas used in the SFE-based test
methods and how can the SFE extraction be used for the
determination of nonpolar TAG and total oil?
How to prepare lipid samples for gas chromatographic
analysis?
What are the functional properties of lipids in bakery prod-
ucts? What are the different roles of nonpolar and polar
lipids in cereal processing?
Which polyunsaturated fatty acids are most common in
plant-based lipids?
Exercises for Readers to Explore the Topic Further
The multiple interactions of lipids with other constituents
(i.e., proteins during dough mixing and starch) are impor-
tant for the processing and quality of cereal foods. How do
different classes of lipids, for example, nonpolar and polar
lipids, influence the functions of gluten proteins and the
starch-gelatinization mechanism?
How can mass spectrometry be used to improve the detec-
tion and identification of lipids?
How can generic breeding increase the nutritional values of
plant-based oils?
See also: Agronomy of Grain Growing: Soybean: Agronomy;
Barley, Rice and Maize Processing: Oil from Rice and Maize;
Breeding of Grains: Soybean: Germplasm, Breeding, and Genetics;
Wheat Breeding: Exploiting and Fixing Genetic Variation by Selection
and Evaluation; Carbohydrates: Starch: Starch Architecture and
Structure; Fats: Healthy Fats and Oils; Grain Marketing and
Grading: Soybean: Grading and Marketing; Oilseed and Legume
Processing: Soybean: Processing; The Basics: Grain: Morphology
of Internal Structure; The Cereal Grains: An Overview of the Family
of Cereal Grains Prominent in World Agriculture; The Legumes and
Pseudocereals: Peanuts: Overview; The Oilseeds: Cottonseed:
Overview; Oilseeds: Overview; Wheat Processing: Analysis of Dough
Rheology in Breadmaking; Baked Product Staling: Mechanisms,
Determinations, and Anti-staling Strategies; Bakeries: The Source of
Our Unique Wheat-Based Food, Bread; Gluten and Modified Gluten;
Ultrastructure of the Wheat Grain, Flour, and Dough.
256 FATS | Lipid Chemistry
Further Reading
AOCS (2000) In: Official Methods and Recommended Practice of the American Oil
Chemists Society. Champaign, IL: The Society.
Barnes PJ (1983) Lipids in Cereal Technology. London: Academic Press.
Christie WW (2010) Lipid Analysis Isolation, Separation, Identification and Structural
Analysis of Lipids, 4th edn. Bridgwater: The Oily Press.
Chung OK (1986) Lipidprotein interactions in wheat-flour, dough, gluten,
and protein-fractions. Cereal Foods World 31: 242244, 246247, 249252,
254256.
Chung OK and Ohm JB (2000a) In: Lorenz KJ and Kulp K (eds.) Handbook of Cereal
Science and Technology, 2nd edn., pp. 417477. New York, NY: Marcel Dekker.
Chung OK and Ohm J-B (2000b) Cereal lipids. In: Karel K and Ponte Jr. JR (eds.)
Handbook of Cereal Science and Technology, 2nd edn. New York, NY: Marcel
Dekker.
Chung OK, Ohm JB, Ram MS, Park SH, and Howitt CA (2009) Wheat Lipids. St. Paul,
MN: American Association of Cereal Chemists Inc (AACC).
Damude HG and Kinney AJ (2008) Engineering oilseeds to produce nutritional fatty
acids. Physiologia Plantarum 132: 110.
Eliasson A-C and Larsson K (1993) Cereals in Breadmaking. A Molecular/Colloidal
Approach. New York, NY: Marcel Dekker.
Gunstone FD (1996) Fatty Acid and Lipid Chemistry. Glasgow: Chapman and Hall.
Hubbard JD, Downing JM, Ram MS, and Chung OK (2004) Lipid extraction from wheat
flour using supercritical fluid extraction. Cereal Chemistry 81: 693698.
King JW (2002) Supercritical fluid technology for lipid extraction, fractionation,
and reactions. In: Lipid Biotechnology, pp. 663687. New York, NY: Marcel Dekker.
King JW (2004) Critical fluid technology for the processing of lipid-related natural
products. Comptes Rendus Chimie 7: 647659.
Macritchie F (1981) Flour lipids Theoretical aspects and functional-properties. Cereal
Chemistry 58: 156158.
Morrison WR (1978a) Cereal lipids. Advances in Cereal Science and Technology
2: 221348.
Morrison WR (1978b) Wheat lipid-composition. Cereal Chemistry 55: 548558.
Morrison WR (1978c) Cereal lipids. In: Pomeranz Y (ed.). Advances in Cereal Science
and Technology, vol. II, pp. 221348. St. Paul, MN: American Association of Cereal
Chemists.
Morrison WR, Tan SL, and Hargin KD (1980) Methods for the quantitative-analysis of
lipids in cereal-grains and similar tissues. Journal of the Science of Food and
Agriculture 31: 329340.
Papantoniou E, Hammond EW, Scriven F, Gordon MH, and Schofield JD (2001)
Isolation of polar lipid classes from wheat flour extracts by preparative high-
performance liquid chromatography. Cereal Chemistry 78: 663665.
Rocha JM, Kalo PJ, Ollilainen V, and Malcata FX (2010) Separation and identification of
neutral cereal lipids by normal phase high-performance liquid chromatography,
using evaporative light-scattering and electrospray mass spectrometry for detection.
Journal of Chromatography A 1217: 30133025.
Salunkhe DK, Chavan JK, Adsule RN, and Kadam SS (1992) World Oilseeds
Chemistry, Technology, and Utilization. New York: Van Nostrand.
Tweeten TN, Wetzel DL, and Chung OK (1981) Physicochemical characterization of
galactosyldiglycerides and their quantitation in wheat-flour lipids by high-
performance liquid-chromatography. Journal of the American Oil Chemists Society
58: 664672.
 Healthy Fats and Oils
SM Ghazani and AG Marangoni, University of Guelph, Guelph, ON, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Healthy minor components in fats and oils.
Fatty acids and triglyceride profiles of different oil sources.
Classification of fats and oils.
Major vegetable oils and beneficial components in them.
Oil refining and its effects on healthy minor components.
Learning Objectives
To understand healthy minor components in oils.
To categorize oils and fats based on fatty acid composition.
To diagnose differences between desirable and undesirable
minor components in oils.
To realize the main purpose of oil refining.
Introduction
Fats and oils are considered an important part of our daily diet
intake because they provide a concentrated source of energy
and essential fatty acids. They are carriers of fat-soluble vita-
mins (A, D, and E) and make fundamental parts of cell walls
and other biological structures. In addition to biological func-
tions and nutritional properties, they exhibit key functionality
by playing an important role in the final quality of food
products. Fats and oils can naturally be found in different
sources, but only a few of these sources are of economic
importance. Oils and fats may mainly originate from plants
(seeds, beans, and fruits), animals (lard, tallow, fish oil, and
milk fat), and microbial sources (yeasts, algal, and bacteria).
Edible oils are mainly obtained from beans, seeds, and fruits by
mechanical and/or solvent extraction. Based on USDA statistic-
al reports, total world vegetable oils production for 2012/2013
was 160.7 million tonnes. Meanwhile, the production levels of
four main vegetable oils in that period were palm oil (55.8
million tonnes) followed with soybean oil (42.8 million
tonnes), canola oil (24.9 million tonnes), and sunflower oil
(13.5 million tonnes). In that period, the total amount of
produced animal fats from four main sources (lard, tallow,
fish oil, and milk fat) was about 25 million tonnes (Table 1).
It is important to consider two important points. First,
some part of produced oils is used for nonedible applications
(e.g., oleochemical industry, biodiesel, cosmetic), and second,
daily dietary intake of fats and oils may go above the total
amount of produced oils because other lipid-containing food
sources such as nuts, meat, and dairy products should be
considered in the diet as a source of fats and oils. Recent studies
in biotechnology and plant molecular biology led to the intro-
duction of numerous types of engineered oilseeds and micro-
bial sources to produce novel fats and oils with new fatty acid
and triglyceride compositions and with a higher oil yield. New
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00100-5
sources are important for their unique quality of healthy minor
components, fatty acid profile, or triacylglycerol composition
(e.g., high oleic oils for frying oil or omega-3-enriched oils). As
fats and oils are the highest source of energy in the human diet
and their main contribution is palatability of many food prod-
ucts, selecting a good source of fats and oils based on the daily
calorie need is important. Choosing an appropriate source of
oil and monitoring the energy intake may be helpful in con-
trolling obesity, which is considered a risk factor for many
chronic diseases such as diabetes, cancer, and cardiovascular
diseases (CVDs).
Health and Nutrition Properties of Fats and Oils
Because fats and oils have different composition when it comes
to health, interest in specifying a fatty acid profile and healthy
minor components has been emerging. Oils and fats are pre-
dominantly a mixture of triacylglycerols as main organic com-
ponents that provide energy (9 kcal g
1) accompanied by
other minor components that could be nutritionally important
such as tocopherols, tocotrienols, sterols, polyphenols,
phospholipids (PLs), sphingolipids, and hydrocarbons. Also,
other minor components are available and mostly removed
during refining steps like waxes, free fatty acids, mono- and di-
acylglycerols, flavoring, and coloring agents.
More than the source of fats and oils, many other factors
such as varieties, growing conditions, maturity of seeds or
fruits, harvesting time, storage conditions, handling, and pro-
cessing settings (extraction and refining oil) may affect the level
of healthy minor components in oils. Based on current recom-
mendations, adults should get 2035% of daily calories from
fats and oil sources. Currently we are emerging with two trends
in food intakefirst, increasing consumption of processed
foods and snacks, and second, increased demand for virgin or
cold-pressed oils. As oils in the snack foods made up a rela-
tively large portion intake of energy in industrialized countries,
choosing snacks could directly affect the health. In this article,
first nutritional properties of healthy minor components will
be discussed followed by a comparison between fatty acid
composition of fats and oils, and main sources of fats and
oils will be reviewed.
Healthy Minor Components
Phospholipids
PLs are organic components with both hydrophilic and hydro-
phobic moieties in molecule structures. They can be found in
all plant and animal cell membranes. The main food sources of
PLs are egg yolk (810%), butter (0.51.2%), and crude vege-
table oils (0.53.7%). The six major types of PLs in vegetable
oils are phosphatidic acid (PA), phosphatidylcholine (PC),
257
258 FATS | Healthy Fats and Oils
Table 1 World main vegetable oils production, 2009/102013/14 (million tonnes)
Oil type/year 2009/10 2010/11
Coconut 3.52 3.70
Cottonseed 4.59 4.96
Olive 3.08 3.25
Palm 46.04 48.73
Palm Kernel 5.60 5.73
Peanut 4.89 5.33
Rapeseed 22.56 23.52
Soybean 38.80 41.30
Sunflower 12.28 12.42
Total 141.36 148.94
a
Forecast, Source: USDA, Foreign Agricultural Service, Oilseeds: World Markets and Trade.
phosphatidylethanolamine (PE), phosphatidyl glycerol, phos-
phatidylserine (PS), and phosphatidylinositol (PI). PLs
showed great positive functionality in several diseases such as
liver diseases, neurological disorders, CVDs, and hepatic and
colon cancers. They also have immunological functionality for
body system, and they can improve physiological and mental
performances. The amount of PLs in crude oils depends on the
oil sources and extraction method. Solvent-extracted oils have
higher phosphatides than mechanically extracted oils.
PLs have been shown to have effects on lowering blood
cholesterol and TAG levels in serum. Many studies have shown
that in athletes, PS can improve performance and decrease
muscle damage during some sports like cycling, weight train-
ing, golf, and endurance running.
Among PLs, PI and PE are more effective than PC in increas-
ing oxidative stability of oils as a synergistic function with
a-tocopherol. In the digestive tract, PLs play an important
functionality in the metabolism of lipids and other digestive
fluids. Although PLs have health-promoting properties, from
an industrial viewpoint, they are considered undesirable com-
ponents. In the neutralization process, they work as an
emulsifier, hinder the separation of oil and water phases, and
help to increase the oil loss in this step. Also, they may interfere
with bleaching clays, poison nickel catalysts throughout hydro-
genation, darkening oil during deodorization and decreased
shelf life of refined oils. PLs can be almost completely removed
from oil through the degumming stage of oil refining; only
they can be found in virgin or cold-pressed oils in different
levels.
Sterols
Sterols comprise of the main part of unsaponifiable matter of
fats and oils. In fats and oils, they can be in two forms of free
and esterified sterols. Free sterols can commonly esterify with
fatty acids, glycosides, and/or ferulic acid to make sterol esters.
The amount of phytosterols in most of common vegetable oils
is usually between 1000 and 5000 ppm (15 g kg
1). How-
ever, higher levels could be found in rapeseed oil
(500011 000 ppm) and in corn oil (800022 000 ppm).
The major phytosterols in common vegetable oils are bras-
sicasterol, campesterol, stigmasterol, b-sitosterol, and avenas-
terol. Canola oil, corn oil, sesame oil, and soybean oil are good
sources of phytosterols between vegetable oils. Some sterol
2011/12 2012/13 2013/14a
3.41 3.65 3.54
5.25 5.27 5.11
3.24 2.67 3.28
51.95 55.77 58.43
6.14 6.51 6.86
5.32 5.55 5.67
24.22 24.90 25.76
42.59 42.81 44.66
15.34 13.53 15.67
157.46 160.66 168.98
esters, for instance g-oryzanol in rice bran oil, have unique
nutritional properties such as antioxidant activity, hypolipi-
demic effects, and stimulation of growth and hypothalamus
activity. Hypocholesteremic properties of phytosterols have
been studied, and based on the guidelines, 2 g of phytosterols
per day, along with 1025 g soluble fibers, was recommended.
In animal fats, the main sterol is cholesterol. Although a
trace level of cholesterol 2050 ppm could be found in vege-
table oils, it is much lower than the levels reported for animal
fats (up to 1000 ppm), fish oils (up to 7000 ppm), dairy fats
(20003000 ppm), and egg yolk (12 500 ppm). It is important
to note that cholesterol is not primarily associated with the
storage adipose (tallow or lard), so lowering the level of total
fat in meat (using lean or extra-lean meats) will not greatly
decrease the cholesterol content because it is a necessary mem-
brane component.
In the human blood system, there are two main types of
cholesterol named high-density lipoprotein (HDL) cholesterol
and low-density lipoprotein (LDL) cholesterol. HDL choles-
terol is good cholesterol because excess HDL cholesterol is
carried to the liver and then metabolized. LDL cholesterol is
bad cholesterol because it remains in the blood plasma and
can start to produce plaques in the arterial wall and lead to
obstruction of heart blood artery, causing a heart attack. So
elevated total and HDL cholesterol levels are considered risk
factors for the development of coronary heart disease that is
one of the main health problem in developed countries. Phy-
tosterols in vegetable oils can inhibit cholesterol absorption.
Intake of phytosterols at the level of 1.53 g day
1 can bring
total blood cholesterol and LDL cholesterol lower by 1015%.
Phytosterols usually have a lower absorption rate in the diges-
tive tract than cholesterol; for example, only 5% of b-sitosterol
of the food supply can be absorbed, so phytosterols in foods
can compete with cholesterol absorption and leads to an
increase in bile secretion and inhibition of cholesterol absorp-
tion from the gut.
Much research has shown that phytosterols, especially
b-sitosterols, inhibited cancer growth and may offer protec-
tion from colon, prostate, and breast cancers.
Besides cholesterol, ergosterol (precursor of vitamin D2)
can also be found in animal fat sources such as fish oil and
butter fat.
Phytosterols are stable molecules, but after as long time
being heated or exposed to oxygen during food processing

and storage, they are susceptible to oxidation, and oxidized
phytosterols (oxyphytosterols) are absorbed from digestive
tract. Oxyphytosterols could be mutagenic and be a source of
free radicals, so they can affect cell viability, stimulate inflam-
mation, cause oxidative changes in the retina, and affect hor-
monal activity. So monitoring the amount of oxidized
phytosterols, especially in long heated or fried foods, would
be necessary.
Tocopherols
The tocols (tocopherol and tocotrienol molecules) are natural
fat-soluble organic components with antioxidant properties.
Many factors may influence the concentration of tocols in
oilseeds such as the amount of unsaturated fatty acids, climate
conditions, and genotype. In extracted oils, the method of oil
extraction has a great influence on tocol concentration in crude
oils. For example, solvent-extracted oils usually contain higher
amounts of tocopherols than cold-pressed oils. In crude oils,
the concentration of tocopherols and tocotrienols varies
between 200 and 1200 ppm. Tocopherols are free or esterified
molecules found in four homologues, namely, a-, b-, g-,
and d-tocopherols. In living organisms (in vivo systems),
a-tocopherol shows the highest vitamin E activity and the
most effective antioxidant properties compared to the other
isomers, whereas in the food system (in vitro) the antioxi-
dant activity of the tocopherol isomers decreases in the
order: d > g > b > a. A single molecule of tocopherol can
react with more than 100 molecules of singlet oxygen, and
this specific property is related to their ability to transform
to their active form after oxidation.
Tocotrienols have more antioxidant activities than tocoph-
erols. They also have health benefits such as hypocholestero-
lemic, antithrombic, and antitumor properties. Tocoterienols,
especially g- and d-, have cholesterol-lowering properties,
probably by inhibiting cholesterol biosynthesis. The recom-
mended dietary allowances daily intake of a-tocopherol (vita-
min E) for adults is 1115 mg. Intake of unsaturated fatty acids
determines the dietary requirement of vitamin E, for example,
for 1 g intake of oleic, linoleic, linolenic acids about 0.1, 0.6,
and 0.9 mg a-tocopherol, respectively, are required.
Both natural and synthetic forms of vitamin E are absorbed
in digestive tract, but the concentration of a natural form in the
blood system was significantly higher than the synthetic form.
Polyphenols
Polyphenols are organic components that mainly influence
flavor and stability of crude oils. At high concentration (more
than 300 ppm), they cause a bitter taste in olive oil. Several
studies have shown that polyphenols have antioxidant and
antimicrobial properties. They showed radical scavenger prop-
erties in the body system, and they may contribute to the lower
incidence of CVD and some types of cancer. Among different
polyphenols, hydroxyltyrosol has the highest antioxidant
properties, and it is more effective than synthetic butylated
hydroxyl toluene.
There are a large variety of polyphenols in crude oils. Virgin
olive oil and cold-pressed canola oils are good sources of
polyphenols.
FATS | Healthy Fats and Oils 259
Hydrocarbons
Saturated and unsaturated hydrocarbons in aliphatic or terpe-
nic forms can be found in fats and oils. Two major well-known
unsaturated terpenic hydrocarbons are carotenoids and squa-
lene. Squalene can be found in olive oil, palm oil, rice bran oil,
and in animal fats (shark liver oil). Squalene is a natural
antioxidant, and it also has antitumor, antibacterial, and
anticarcinoma properties.
Carotenoids are yellowish to red color natural family pig-
ments found in most oils. b-Carotene (pro-vitamin A) and
lycopene are good examples of carotenoids. b-Carotene
exhibits vitamin A activity and, along with other carotenoids,
acts as a strong antioxidant in the presence of light. Research
showed that b-carotene and a-tocopherol have a synergistic
antioxidant effect.
Red palm oil is the richest source of carotenoids, whereas
butter and cod liver oil have large amounts of carotenoids in
animal fats. Research showed that a high level of carotenoids in
red palm oil has an anticarcinogenic effect.
Vitamin A deficiency is common worldwide and leads to
eye diseases, severe infection, and death in many people, espe-
cially children and women. The body can make vitamin A from
carotenoids in oils. The recommended daily intake of vitamin
A (retinol) for females is 700 mg and for males 900 mg. The
main functionalities of vitamin A are in visual system, growth,
immune system, epithelial cellular integrity, and reproduction.
Fatty Acids and Triacylglycerols Compositions
More than 1000 types of fatty acids have been identified.
However, not all of them are important from nutrition and
health points of view. The main differences between fatty acids
are the number of carbon atoms and the number and position
of double bonds. Fatty acid double bonds occur in two geo-
metric isomers either in the cis form that is the natural form of
fatty acids (hydrogens on the same side of the plane), or trans
configuration (hydrogens on opposite sides of the plane),
which form during hydrogenation or high temperature expo-
sure during deodorization. Some fatty acids have special func-
tional groups such as hydroxyl, epoxy, keto, or ring groups. The
fatty acids are classified as short-chain fatty acids (C2C4),
medium-chain fatty acids, (C6C10), and long-chain fatty
acids (C12C24) and saturated (without double bonds),
mono-unsaturated (one cis or trans double bond), and poly-
unsaturated (two and more cis or trans double bonds).
Numbering of fatty acids could be based on the omega-
reference system that indicates the number of carbons, the
number of double bonds, and the position of the double
bond for the carbon atom of x, starting from the terminal
methyl group and designated as (n
x) for example C18:3
(n
3). The main saturated fatty acids are butyric acid, C4:0;
caproic acid, C6:0; caprylic acid, C8:0; capric acid, C10:0; lauric
acid, C12:0; myristic acid, C14:0; palmitic acid, C16:0; stearic
acid, C18:0; arachidic acid, C20:0; behenic acid, C22:0; and
lignoceric acid, C24:0. The main monoenoic fatty acids are
lauroleic acid, C12:1 (n
3); myristoleic acid, C14:1 (n
5);
palmitoleic acid, C16:1 (n
7); oleic acid, C18:1 (n
9); gado-
leic acid, C20:1 (n
11); cetoleic acid, C22:1 (n
13); and
erucic acid, C22:1 (n
9).
260 FATS | Healthy Fats and Oils
-Carotene
Squalene
HO
H3C O
CH3
-Tocopherol
HO
Cholesterol
Polyunsaturated fatty acids (PUFAs) can be characterized
based on a methylene group that can be central, conjugated, or
unconjugated. For instance, linolenic acid (C18:3) can be
found in two forms of a-linolenic -9, 12, 15-oic acid, (n
3)
or g-linolenic-6, 9, 12-oic acid (n
6). Eicosapentaenoic acid
(EPA) C20:5 (n
3) and docosahexenoic acid (DHA) C22:6
(n
3) found in oily fishes are examples of PUFAs with nutri-
tional characteristics.
Triacylglycerols comprise more than 95% of fats and oils
composition. Each triacylglycerol molecule consists of three
fatty acids that attached to the glycerol molecule with ester
bonds. Normally triacyglycerols can categorize to simple tri-
glycerides with only one type of esterified fatty acid, for
example, triolein (OOO); di-acid triglycerides with two types
of fatty acids esterified to glycerol molecule, for example, di-
palmito 1,3 olein 2 (POP); and tri-acid with three types of acyl
groups, for example, palmito 1, oleo 2, and stearin 3 (POS).
Glycerol molecules can also esterify with only one fatty acid to
form monoacyl glycerols or with two same or different fatty
acids to make diacylglycerols. The physical, chemical,
biological, and physiological properties of fats and oils related
to triacylglycerol composition and presence of healthy minor
components. The reason for the importance of triacylglycerol
composition is that two oils with the same fatty acid profile
may have different physical, chemical, and physiological prop-
erties. Also the position of fatty acids on glycerol molecules has
health and physiologically importance. For instance during fat
digestion, because human lipase is specific (hydrolyzing only
fatty acids in positions 1 and 3 of TAGs), the fatty acids
attached to carbon number 2 of glycerol structure (the sn-2
position) are especially absorbed as the 2-monoacylglycerol
OH
O
O
Sesamol
HO
-Sitosterol
and serve as the template for re-esterification by intestinal
cells to reform triacylglycerols. In vegetable oils, unsaturated
fatty acids are esterified in the sn-2 position, whereas in animal
fats saturated fatty acids are in this position. In addition, selec-
tive distribution of fatty acids on glycerol backbone is an
important factor that leads to triacylglycerols diversity. For
example, palm oil contains six types of fatty acids, but only
14 diverse triacylglycerols can be detected.
Omega-3 and omega-6 PUFAs have different physiological
properties in the human body system. Whereas n
3 fatty acids
have anti-inflammatory, antithrombotic, antihypertensive,
and anti-arrhythmic derivatives, n
6 fatty acids cause inflam-
matory, thrombotic, hypertensive, and arrhythmic metabo-
lites. That is the reason for the importance of the n
6/n
3
ratio balance in human diet.
The impact of saturated fatty acids on LDL cholesterol
depends on the number of carbon atoms in the fatty acid
chain with lauric acid > myristic acid > palmitic acid > stearic
acid. Clinical studies showed that saturated fatty acids lead to
an increase in total blood cholesterol and LDL cholesterol,
whereas mono-unsaturated and more effective
polyunsaturated fatty acids lead to a decrease in the total
serum cholesterol and LDL cholesterol and increasing HDL
cholesterol in the bloodstream. The risk of coronary heart
disease in the societies in which people consume olive or
canola oil (good source of oleic acid) and fish oil (good source
of omega-3 fatty acids) is lower.
Studies have shown that consumption of n
3 fatty acids
(fish oil, walnut oil, and flaxseed oil) reduces mortalities
related to CVDs by lowering triacylglycerol accumulation in
the liver and circulation.

Classification of Oils and Fats Based on Fatty Acid
Profile
Medium-Chain Fatty Acids (C6:0 to C10:0) Subgroup
This group of oils (MCTs) is important in the case of providing
an instant source of energy for the body. These fatty acids have
small size and higher solubility and are mainly transported
directly via portal circulation and metabolized in the liver,
whereas longer-chain fatty acids are absorbed via the lymphatic
system after micellar transport at the intestinal wall. MCTs are
neutral on LDL-, HDL-cholesterol or triacylglycerol concentra-
tion in serum and may be beneficial in controlling obesity.
Lauric Acid and Myristic Acid Subgroup
Among saturated fatty acids, lauric acid and myristic acid are
the most blood cholesterol and LDL-cholesterol increasing
fatty acids. Although, the effect of myristic acid is higher than
lauric acid, the position of fatty acids on the glycerol backbone
needs to be considered. Coconut oil and palm kernel oil are
categorized in this subgroup.
Palmitic Acid Subgroup
The effect of palmitic acid on increasing serum cholesterol and
LDL cholesterol is lower than lauric acid and myristic acid.
Palm oil and lard are good examples of this group.
Stearic Acid Subgroup
Among saturated fatty acids, stearic acid exhibited as quite
neutral with no deleterious effect on plasma lipids. Cocoa
butter and shea butter can be in this subgroup because they
contain up to 40% of stearic acid.
Oleic Acid Subgroup (>60%)
Olive oil is the best representative oil of this subgroup, which
contains about 80% of oleic acid and then canola oil with
about 65% oleic acids. In nuts oil, hazelnut oil and macadamia
oil are rich in oleic acid.
Linoleic Acid Subclass (>60%)
The common vegetable oils gathered in this group are grape
seed oil, evening primrose oil, and safflower oil. As the human
body cannot synthesize linoleic acid from food components,
presence of this fatty acid is essential for good health and
termed an essential fatty acid. As mentioned earlier, the inflam-
matory effect of consuming n
6 fatty acids should be consid-
ered and n
6/n
3 ratio should be balanced in the diet.
Linoleic Acid Saturated Fatty Acids Subgroup
Cottonseed oil and watermelon seed oil are clustered in this
group. Their fatty acid profiles mainly contain linoleic acid,
associated with saturated fatty acids (especially palmitic acid).
FATS | Healthy Fats and Oils 261
Linoleic Acid Oleic Acid Subgroup
Sunflower oil, corn oil, sesame oil, pumpkin seed oil, and
borage oil are good examples of this subgroup. In this group,
borage oil has a high amount of g-linoleic acid (about 22% of
the total fatty acids).
Conjugated Linoleic Acid
Conjugated linoleic acid (CLA) is a term used for positional
and geometric isomers of linoleic acid that can be found in
animal fats such as beef, lamb, and dairy foods. CLA are
produced by microbial fermentation of PUFAs and isomeriza-
tion of linoleic acid in the rumens of ruminants. Whereas
double bonds in linoleic acid are between the ninth and
tenth carbons and the twelfth and thirteenth carbons, CLA
has conjugated double bonds at carbon atoms 10 and 12 or 9
and 11, with possible cis and trans combinations. Many clin-
ical studies have shown that conjugated linoleic acid may have
physiological effects, including anticarcinogenic, anti-
atherogenic, improvement of type II diabetes, and immuno-
modulating properties of CLA.
Some clinical studies on animals showed that conjugated
linoleic acid is quite effective to inhibit the growth and metas-
tasis of breast and prostate cancers.
Trans Fatty Acids
Two types of trans fatty acid (TFAs) isomers can be found in
fats as naturally and industrially produced fatty acids. The
presence of TFAs in animal fats was revealed some time ago.
TFA in tallow and milk fat is vaccinic acid C18:1 (n
7t). This
TFA is naturally produced during biohydrogenation and isom-
erization of PUFAs in rumen guts.
Catalytic hydrogenation is an industrial modification pro-
cess that is applied to reduce the amount of polyunsaturated
fatty acids and leads to improvement in oxidative stability of
oils and increasing hardness. This industrial process produces
TFA, which are almost unavailable in natural fats. TFA levels
and isomeric distribution depend on processing factors (nickel
catalyst type and concentration, hydrogen pressure, stirring
speed, and reaction temperature), fatty acid composition of
original oil, and degree of saturation (partially or fully
hydrogenation).
Partial hydrogenation produces mostly geometrical and
positional trans-18:1 isomers (mainly elaidic acid) while
reducing the amount of PUFA. During deodorization of edible
oils, a small amount of trans-linoleic and trans-linolenic acid
are produced.
A positive correlation between intake of a high amount of
TFAs in a diet with increased risk of chronic diseases such as
cardiovascular, inflammatory, autoimmune diseases, and can-
cer have been found. TFAs raise the amount of LDL cholesterol
and lower the level of HDL cholesterol, thus resulting in dou-
bly detrimental effects. New regulations on TFA labeling,
which led to a ban on industrially produced trans fats in
foods, caused a dramatic reduction in TFAs in food products,
but they still can be found in some foods such as confectionery
coatings and fillings, biscuits (cookies), pie crusts, toffees, and
analogs of dairy products such as butter, cream, and toppings.
262 FATS | Healthy Fats and Oils

Using naturally saturated fats such as palm oil and/or
applying fully hydrogenated oils or vegetable oils from new
varieties such as high oleic oils could be solutions for food
industries to substitute TFAs in their products.
Omega-3 Fatty Acids
The omega-3 fatty acids consist of a-linolenic acid, EPA, and
DHA. Several clinical studies showed a positive correlation
between consumption of omega-3 fatty acids in the diet and
an improved prognosis of CVD and nonalcoholic fatty liver
disease. The main sources of a-linolenic are flaxseed oil and
walnut oil and oily fishes such as tuna, menhaden, anchovies,
sardines, and mackerel are the main sources of EPA and DHA.
Major Vegetable Oils
This section is mainly focused on the main sources of vegetable
oils that would cover more than 95% of total vegetable oils
consumed in the word (Table 1).
Palm Oil
Palm oil is the main vegetable oil produced in the world.
During 20132014 about 53.4 million tonnes of palm oil
were produced (Table 1). Palm oil originated from South Africa
and was introduced to East Asia in 1884. Currently, Indonesia
and Malaysia are the two main palm oil producers in the world.
Palm oil is rich in healthy minor components such as carbohy-
drates (carotenoids and squalene) and tocols. Crude palm oil
has about 4001000 ppm carotenoids (pro-vitamin A) but
after physical refining, the amount of carotenoids decreased
to nondetectable levels. The squalene content in crude palm
oil is between 200 and 500 ppm, which is dramatically higher
than other common vegetable oil sources. Total tocols in crude
palm oil are about 800 ppm, 70% of which are tocotrienols.
The distribution of tocopherols in palm oil is a-tocopherol
22%, and b-, g-, and d- tocopherols represent 20%, 46%, and
12%, respectively. Crude palm oil contains 210620 ppm of
phytosterols. The main phytosterols in palm oil are b-sitosterol
(120370 ppm), campesterol (46150 ppm), and stigmasterol
(2666 ppm).
Palm oil is a good source of carotenoids. Based on the
carotenoid content, crude palm oil color varies from yellowish
to dark red. The main carotenoids in palm oil are a-carotene
(2442%) and b-carotene (5060%). Palmitic acid and oleic
acid are the main fatty acids in palm oil (more than 80% of
total fatty acids and almost in equal amount). Normally, palm
oil is fractionated into different derivatives (palm olein, palm
stearin, palm super olein, and palm mid fraction) with specific
physical and chemical characteristics (Tables 2 and 3). Because
of fatty acid composition (about 50% saturated fatty acids and
50% oleic acid), crystallization in the form of b0 and the

Table 2 Fatty acid composition of palm oil and its derivatives

Fatty acids (%) Palm oil Palm olein Palm super olein Palm top olein Palm stearin Palm mid fraction

12:0 0.2 0.3 0.3 0.2 0.10.3 0.00.3

14:0 1.1 1.1 1.0 1.0 1.11.7 0.81.4
16:0 44.1 40.9 35.4 28.8 49.868.1 41.455.5
16:1 0.2 0.2 0.3 0.3 0.050.1
18:0 4.4 4.2 3.8 2.5 3.95.6 4.76.7
18:1 39.0 41.5 45.1 52.0 20.434.4 32.041.2
18:2 10.6 11.6 13.4 14.6 5.08.9 3.611.5
18:3 0.3 0.4 0.3 0.4 0.10.5 0.00.2
20:0 0.2 0.4 0.3 0.2 0.30.6 0.00.6

Malaysian Palm Oil Board (MPOB) (2009) Pocketbook of Palm Oil Uses, pp. 341.

Table 3 Fatty acid composition of main vegetable oils

Fatty acids (%) Soybean oil Canola oil Sunflower oil Cottonseed oil Corn oil Peanut oil Olive oil

12:0
14:0 0.1 0.5-1.3 <0.1 <0.1
16:0 11.0 4.3 57 1731 813 813 7.520
16:1 0.1 0.2 <0.4 <1 <1 <0.3 0.33.5
18:0 4.0 1.8 46 13 14 34 0.55
18:1 23.4 62.3 1525 1321 2432 4866 5583
18:2 53.2 19.4 6270 3460 5562 1428 3.521
18:3 7.8 9.2 <0.2 <1 <2 <0.3 <1
20:0 0.3 0.6 <1 0.20.5 <1 12 <0.8
20:1 1.2 <0.5 <0.5 12
20:2 0.1
22:0 0.1 <1 <0.5 24 <0.3
24:0 12 <1

presence of natural antioxidants (tocols), this oil is widely used
in the food industry as a main part of base stock to formulate
margarines, shortenings, frying oils, and confectionary fats.
Soybean Oil
Soybean is the main oilseed produced in the world, and soy-
bean oil is the second-largest-produced vegetable oil after palm
oil (Table 1). The worlds major producers of soybean oil are
the United States, China, Argentina, Brazil, EU-27, and India.
Crude soybean oil is a good source of phosphatides that after
extraction and purification steps can be used as a natural
emulsifier (lecithin) in various food products. The three
major phosphatides in crude soybean oil are PC, PE, and PI
present in the ratio of 55%, 26%, and 18%, respectively. Con-
centration of phytosterols and tocopherols in soybean oil is
23504050 ppm and 12571370 ppm, respectively. In soy-
bean oil, b-sitosterol (4759%) and g-tocopherol (4460%)
are the major phytosterol and tocopherol in soybean oil. Soy-
bean oil contains about 68% a-linolenic acid. Research
showed that soybean oil consumption led to a reduction in
total blood serum cholesterol and LDL cholesterol.
The major triacylglycerols in soybean oil are OLL (20%),
LLL (19%), and PLL (13%). Crude solvent extracted soybean
oil typically has about 2%PLs. The main phytosterols in soy-
bean oil are b-sitosterol (52%), campesterol (25%), and stig-
masterol (23%). Two major hydrocarbons in soybean oil are
squalene and hexahydrofarnesyl acetone.
Canola Oil
Canola oil was first produced in Canada in 1974. Genetic
modifications of rapeseed oil led to a reduction in the concen-
tration of erucic acid and glucosinolate. The currently rape-
seeds in use have erucic acid and glucosinolates less than
0.1% and 8.5 mM g
1, respectively. After palm oil and soybean
oil, canola oil is the third major vegetable oil produced in the
world (Table 1), and China, EU-27, Canada, India, and Japan
are the main producers. Canola oil in Canada is extracted
mainly from the seeds of genetically modified Brassica napus
L., but a small part of extracted canola oils are from Brassica
rapa L. Canola oil is the only known vegetable oil with a sulfur
atom in some fatty acid structures that are responsible for the
sulfur flavor in the oil.
Many dietitians believe that canola oil could be considered
the healthiest edible oil. Canola oil has unique characteristics
such as fatty acid composition and levels of tocopherols,
phytosterols, and polyphenols. Canola oil contains about
12% a-linolenic acid (omega-3) and about 65% oleic acids.
Also, it contains a low amount of saturated fatty acids (<7%)
compared to other common vegetable oils.
This oil has a high amount of polyphenols (113 ppm gallic
acid), and the predominant polyphenols in crude canola oil
are vinylsyringol, 4-vinylsyringol dimer, sinapic acid, and
sinapine.
PA is the main phospholipid in crude canola oil (around
44%). Canola oil lecithin is usually used as an animal feed
ingredient. After soybean oil and corn oil, canola oil has the
highest amount of tocopherol isomers, whereas the a- and
g-tocopherols are the major tocopherols with a 1:2 ratio
FATS | Healthy Fats and Oils 263
(100386 to 189753 ppm, respectively; data adapted from
Codex Alimentarius, 2010). Green color of crude canola oil is
related to the presence of chlorophylls (550 ppm).
Plant breeders have developed canola oil with new fatty acid
composition such as high oleic canola oil (oleic acid raised to
about 85%), high lauric canola oil (up to 31% lauric acid), and
high stearic acid canola oil (stearic acid content increased to
about 28%).
Sunflower Oil
Russia, Ukraine, Turkey, European countries, and Argentina are
the main producers of sunflower oil. During 20132014,
about 15.7 million tonnes of sunflower oil were produced
(Table 1). Linoleic acid as an essential fatty acid (6270%)
and oleic acid (1525%) are predominant fatty acids in this
oil. These amounts for new mutant of high oleic sunflower oil
are 217% and 7591%, respectively. The main triglycerides in
regular sunflower oil are LLL (36%) followed by OLL (29%).
Sunflower oil normally has about 32005100 ppm phytos-
terols (5864% b-sitosterol) and 4401200 ppm tocopherols
(9197% a-tocopherol). It is also considered a good source of
squalene (150200 ppm). Crude sunflower oil has 0.51.2%
PLs and around 180 ppm of total polyphenols. The main poly-
phenols in crude sunflower oil are caffeic acid, vanillic acid,
and chlorogenic acid.
Cottonseed Oil
Cottonseed oil extraction plants are mainly in China, India,
Pakistan, the United States, Brazil, and Turkey (5 million
tonnes per year production; Table 1). Typically the three
main fatty acids in this oil are palmitic acid, oleic acid, and
linoleic acid with an average percentage of 22, 20, and 54,
respectively. Four major triacylglycerols in cottonseed oil are
PLL (27.5%), LLL (19%), POL (14%), and OLL (12.5%). Cot-
tonseed oil has unique types of fatty acidsmalvalic and ster-
culic acids, which are called cyclopropenoid fatty acids. Both
malvalic and sterculic acids have one double bound at the
propene ring and some toxicity properties, but after deodori-
zation the amount of total cyclopropenoid fatty acids will
reduce to 0.04%. Typically, PLs content in crude cottonseed
oil is about 0.70.9%. Cottonseed oil is also exceptional for the
presence of a toxic polyphenolic component named gossypol.
This pigment gives a dark red color to crude cottonseed oil.
Most parts of the gossypol are removed during neutralization,
and at the end of refining a safe level of 15 ppm would be
present in oil. Although gossypol shows toxicity effects, recent
research showed that this component has bioactive properties
such as antimicrobial, antioxidant, and anticancer activities.
b-Sitosterol is the most abundant sterol in cottonseed oil fol-
lowed by campesterol, stigmasterol, and D5-avenesterol. Cotton-
seed oil is a rich source of tocopherols. The total tocopherol
content in crude cottonseed oil is about 1000 ppm, and a- and
g-tocopherols account for 41% and 58%, respectively. The pres-
ence of a high level of natural antioxidants, a relatively high
percentage of saturated fatty acids, and a special nutty flavor
makes this oil one of the best choices to use as a frying oil.
264 FATS | Healthy Fats and Oils
Corn Oil
Corn oil is a by-product of corn meal- and starch-making
companies. Fatty acid composition of corn oil comprises a
relatively high level of linoleic acid (5862%). The major
triacylglycerol molecules in corn oil are LLL (25%), LLO
(22%), LLP (15%), OOL (11%), and PLO (10%).
In vegetable oils, corn oil is categorized as one of the richest
source of phytosterols (830025 500 ppm) and tocopherols
(11301830 ppm). The main phytosterol and tocopherol in
corn oil are b-sitosterol (6370%) and g-tocopherol
(6889%), respectively. Numerous clinical studies showed a
correlation between corn oil consumption and total serum
cholesterol reduction that may be related to a high concentra-
tion of phytosterols in this oil. Research showed that a high
ratio of g-tocopherol in corn oil has health benefits such as a
protection effect against DNA damage, blood pressure, platelet
aggregation, and diabetes.
Peanut Oil
China and India are the main producers of peanut oil. Phos-
pholipid content of crude peanut oil is 0.62%, depending on
the maturity of peanuts. The main fatty acids in peanut oil are
oleic acid (4553%), linoleic acid (2732%), and palmitic
acid (1114%). This oil specifically has long-chain saturated
fatty acids such as arachidic acid (12%), behenic acid
(1.54.5%), and lignoceric acid (0.52.5%). The main triacyl-
glycerols in peanut oil are OOL (19%), OLL (18%), POL
(13%), OOO (12%), and POO (7%). Crude peanut oil con-
tains about 3400 ppm total phytosterols, whereas the main
phytosterols are b-sitosterol (64%), campesterol (15%), stig-
masterol (11%), and D5-avenesterol (8%). Tocopherols con-
centration in peanut oil ranges from 130 to1300 ppm with the
composition of a-tocopherol (4765%) and g-tocopherol
(3050%). This oil also has a high level of squalene
(400700 ppm).
Olive Oil
Olive oil has been used for a long time as a food and a
medicine. The world average annual production of olive oil is
about 3 million tonnes mainly in Mediterranean countries.
Many epidemiological studies have shown that the lower
rates of breast, ovarian, and colorectal cancers in Mediterra-
nean countries may be related to consumption of olive oil.
There are two kinds of olive oil in the world: one with a high
amount of oleic acid and low amount of linoleic acid
(European origin) and another with the opposite ratios
(North African origin). This diversity caused a wide range of
fatty acid and triacylglycerol composition to be considered for
this oil. In general, three major fatty acids in olive oil are oleic
acid (5583%), palmitic acid (7.520%), and linoleic acid
(3.521%). The greenish color of virgin olive oil is related to
the presence of chlorophyll in this oil, and their level ranges
from 10 to 30 ppm. a-Tocopherol is the major tocopherol (up
to 95%) in olive oil, and it has a range from 100 to 300 ppm.
The level of squalene in olive oil is relatively high and in most
of samples reported in the range of 200500 ppm. Research
showed that squalene in olive oil may have a preventive effect
on breast cancer, and it also has cardioprotective properties.
Total phytosterol content in virgin olive oil is between 980
and 1840 ppm; meanwhile the main phytosterol is b-sitosterol
(7590%). Virgin olive oil is considered a rich source of poly-
phenols. The amount of polyphenols in virgin olive oil
changes from 100 to 300 ppm. Polyphenols in virgin olive
oil (tyrosol, hydroxytyrosol, and oleuropein) have very strong
free radical scavenger properties and antioxidant activity.
Research showed that they may correlate with decreasing the
risk of CVDs and cancer. The level of phosphatides in olive oil
is lower than common seed oils (40135 ppm). Clinical stud-
ies have shown that olive oil consumption has lots of preven-
tive effects against chronic diseases such as obesity, CVDs,
inflammatory diseases, and cancer.
Coconut Oil
Average annual production of coconut oil is 34 million
tonnes, mainly in the Philippines, Indonesia, and India. Nor-
mally, coconut oil is extracted after sun or fire drying of the
coconut kernel. Dried kernel is named copra, and it has about
65% oil. Coconut oil contains high levels of saturated fatty
acids mainly lauric acid and myristic acid (more than 60% of
total fatty acids). Coconut oil and palm kernel oil are the
richest natural source of medium-chain fatty acids. Because
the major fatty acids in coconut oil are saturated, the total
tocopherol level in this oil is very low (55 ppm). The main
fatty acid and triacylglycerol in coconut oil are lauric acid
(48%) and trilaurin (19%), respectively. The special flavor of
coconut oil is related to the presence of d- and g- lactones.
The range of total phytosterol in this oil is between 800 and
2300 ppm. The major sterols are b-sitosterol (45%), D5-
avenasterol (30%), and stigmasterol (14%).
The amount of hydrocarbons in coconut oil is high,
whereas it only has a small amount of squalene (20 ppm),
but the level of polycyclic aromatic hydrocarbons (PAHs)
could reach to about 3000 ppm as result of contamination of
oil with petroleum oil or during fire drying. The main differ-
ence between fatty acid composition of coconut oil and palm
kernel oil is that although the level of medium-chain fatty acids
(caprylic and capric acids) in coconut oil is higher than palm
kernel oil, the amount of oleic acid in palm kernel oil is higher
than coconut oil. These oils have a sharp melting point that
makes them highly suitable to formulate different food prod-
ucts such as confectionary fats, ice creams, and icings (Table 4).
Sesame Oil
For a long time, sesame was used as an oilseed crop because of
high levels of oil (50%) and protein (2025%) in this seed.
Sesame oil is produced mainly in China, Japan, India,
Singapore, and Mexico. In sesame oil, oleic acid and linoleic
acid are the main fatty acids with the same ratio (around 40%).
The major triacylglycerols in this oil are LLO (25%), LLL
(20%), and LOO (15%). Total phytosterols and tocoph-
erols contents in sesame oil are 54006300 ppm and
200500 ppm, respectively. b-Sitosterol and campesterol are
the main phytosterols in this oil (more than 80% of total
sterols), and g-tocopherol comprises more than 96% of

Table 4 Fatty acid composition of lauric oils
FA (%) Palm Kernel
6:0 <0.8
8:0 25
10:0 35
12:0 4451
14:0 1517
16:0 710
16:1 <0.1
18:0 23
18:1 1218
18:2 14
18:3 <0.7
20:0 <0.3
20:1 <0.5
Karkeskind A (1996) Sources of the main fats and relevant monographs. In: Oils and
Fats Manual, pp. 117323. Paris: Lavoisier Publishing.
total tocopherols. Sesame oil has a good oxidative stability.
More than tocopherols, the stability of this oil is related to
the presence of the unique natural phenolic antioxidants of
sesamol and sesamolin (totally around 5000 ppm). In
many countries, sesame oil is used as frying oil because
of its high stability and also its nutty flavor.
Rice Bran Oil
Rice bran oil is mainly produced in Japan, Thailand, India,
China, and Vietnam. The major fatty acids in rice bran oil are
oleic acid, linoleic acid, and palmitic acids (typically, 42%,
32%, and 20%, respectively). Activity of lipase during rice
bran oil extraction leads to an increase in the amount of FFA,
mono-, and diacylglycerols. Triglycerides totally comprise
8386% of total neutral lipids, whereas PLO, PLL, and OOO
are predominant triacylglycerols in this oil. The concentration
of phytosterols in rice bran oil is higher than common
vegetable oil (32 000 ppm) with the composition of 54% of
b-sitosterol, 20% of campesterol, and 11% of D5-avenesterol.
Rice bran oil is a good source of tocopherols, and the average
of tocopherol content in rice bran oil is 860 ppm. Research has
shown that rice bran oil has health benefits such as lowering
serum cholesterol level, inhibiting platelet aggregation,
increasing fecal bile excretion, and decreasing early
atherosclerosis.
Nut Oils
The main worlds most popular nuts are peanut, almond,
cashew, pistachio, walnut, hazelnut, and pecan. The oil content
of nuts varies from 0.8% for chestnuts to 67% for Brazil nuts.
Fatty acid composition of nut oils is very diverse. The highest
amount of saturated fats belongs to Brazil nut oil (around 23%),
and the lowest level of saturated fat belongs to heartnut oil
(about 5%). Hazelnut oil and pistachio oil have the highest
amount of oleic acid (more than 75%), and walnut oil has the
highest amount of n
3 and n
6 PUFAs. a- and g-Tocopherols
are the main tocopherols in nut oils. a-Tocopherol is the
predominant tocol in hazelnut, pistachio, almond, and peanut
FATS | Healthy Fats and Oils 265
oils, but g-tocopherols is the major tocopherol in chestnut,
pecan, walnut, heartnut, cashew, and Brazil nut oils. Macadamia
Coconut nut oil is rich in tocoterienols (about 60 ppm). Walnut oil has
<0.1 the highest amount of total phytosterols (3000 ppm) followed
610 by peanut and pecan oils (2800 ppm) and almond oil
510 (2700 ppm). Macadamia oil has the lowest level of phytosterols
3954 (1280 ppm).
1523 Based on clinical studies, regular nut consumption or using
611 nut oils as salad oil is associated with reduced risk of coronary
<2 heart disease, diabetes, and cancer.
14
411
12
Fats and Oils Refining
<0.1
<0.2 The main purpose of crude oil refining is to remove undesir-
<0.2 able minor components and contaminants from oils. Free fatty
acids, oxidation products, chlorophyll, waxes, and trace metals
from oils are generally undesirable because they may decrease
the quality of refined oil through darkening, foaming, smok-
ing, precipitation, development of off-flavor, and decreasing
oxidative stability or be potentially dangerous to health. The
source of oil contamination could be from oilseeds during
harvesting and storage with trace metals, dioxins and poly-
chlorobiphenyls from industrial wastes, polycyclic aromatic
hydrocarbons, or mineral oil from vehicle exhausts. Another
source of oil contaminants could be the trace amount of insec-
ticides, fungicides, or herbicide extracted from oilseeds and
transferred to oils. Although most of these chemicals are
removed during refining, oil refining may also lead to the
formation of another group of degraded products such as
polycyclic aromatic hydrocarbons (PAHs) or 3-monochloro-
propane-1,2-diol (3-MCPD) esters and glycidolesters.
Another important fact about oil refining is related to the
loss of micronutrients during refining. Fully refining (degum-
ming, neutralization, bleaching, and deodorization) of oils
leads to change not only in the level of healthy components
but also in their chemical structure and composition by isom-
erization or polymerization. Research showed that traditional
refining leads to about a 20% loss in total amount of tocoph-
erols and 24% in phytosterols level in canola oil.
A typical specification for fully refined oil in general could
be as taste (bland), moisture (max. 0.05%), phosphorus (max.
5 ppm), free fatty acids (max. 0.1%), peroxides (max.
3 meq/kg), iron (max. 0.5 ppm), copper (0.05 ppm), lead
(max. 0.01 ppm), hexane (max. 5 ppm), benzopyrene (max.
2 ppb), dioxins (0.75 pg), aflatoxins (6 ppb), and mineral oil
(LOD).
Conclusion
Although fats and oils contribute to palatability of many food
products, consumers are nowadays paying more and more
attention to their health and well-being. Based on WHO rec-
ommendation for dietary intake of fats and oils for adults, it is
suggested that we get 2035% of our energy from fats and oils
sources. In a selection of healthy oil sources, many factors
should be considered such as fatty acid composition, healthy
minor components, process of oil extracted (solvent extracted,
hot-pressed or cold-pressed), and minimally refined or fully
266 FATS | Healthy Fats and Oils
refined oil. Other factors such as availability, price, food appli-
cation, and stability should be considered.
It is also important to know that in the food industry oil or
fat used in a food product has been carefully chosen for its
unique physical or functional characteristics that are not
always a healthy one. This has been a challenge for food
companies to offer a healthy food with a delicious taste.
Exercises for Revision
What makes an oil or fat healthy?
Canola oil is a healthy oil, but is it suitable for deep-frying
foods?
What is the main difference between essential fatty acids
and omega-3 fatty acids?
Is it true to say cold-pressed oils have higher levels of
healthy minor components compared to solvent-extracted
oils?
Exercises for Readers to Explore the Topic Further
Microbial fats and oils sources and their health benefits.
New genetically modified oils.
Structured lipids and their health functionality.
See also: Barley, Rice and Maize Processing: Maize: Wet
Milling; Oil from Rice and Maize; Oilseed and Legume
Processing: Soybean: Processing; The Oilseeds: Canola: Overview;
Oilseeds: Overview; Soybean: Overview.
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 Bioactives and Toxins
Contents
Bioactive Compounds in Wheat Bran
Bioactives: Antioxidants
The Antinutritional Components of Grains
Mycotoxins
Bioactive Compounds in Wheat Bran
HD Sapirstein, University of Manitoba, Winnipeg, MB, Canada
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
The structure of the wheat kernel is described.
There is an important difference between wheat bran as it
exists in the seed, which can be hand-dissected, and wheat
bran commonly obtained by milling.
Food-grade bran is not a standardized product and can vary
in purity and composition of bioactive compounds due to
genetic factors (different wheat varieties or types), the
wheat growing environment, and also how the milling is
carried out.
The unique protective role of bran from a seeds perspective
reflects its complement of bioactive compounds, particu-
larly those in fiber tissues.
Bran is the most complex physical component of the wheat
kernel containing many layers of tissues with more or less
distinct composition of bioactive compounds, particularly
between the outer pericarp and inner aleurone.
The aleurone layer of bran has a high concentration of the
fiber compound arabinoxylan and many other bioactive
phytochemical compounds such as minerals, B vitamins,
phytic acid, and ferulic acid.
Bran is a heterogeneous mixture of several different types of
nonstarch polysaccharides, which together constitute the
fiber fraction of bran that represents most of the fiber in
whole wheat.
The typical contents of bioactive compounds that are sig-
nificantly enhanced in bran compared to levels found in
whole wheat are presented.
Learning Objectives
To recognize and understand the diversity of bioactive
compounds that are contained in wheat bran, along with
their heterogeneous distribution in the different bran tissue
layers, and the degree to which these bioactive compounds
are enhanced compared to flour or whole wheat.
268 Encyclopedia of Food Grains, Second Edition
Introduction
Wheat, eaten as bread and other baked products, has been a staple
of the human diet for several thousand years. Wheat is a globally
important agricultural commodity and ubiquitous food ingredi-
ent mainly because it is the only cereal grain capable of being
processed into leavened bread, an attribute that derives from its
unique gluten protein composition. While the use of whole
wheat for food waned significantly in the early part of the twen-
tieth century in favor of baked goods made with refined flour, the
past several decades have seen a remarkable resurgence in the
consumption of whole wheat and whole grains in general. This
has been sparked by increasing consumer interest in healthful
eating and awareness of the importance of the health-protecting
and health-promoting effects of whole grains. The functional
properties of whole wheat for health extend well beyond its
nutritional properties and derive for the most part from bioactive
compounds in the bran fraction. Those compounds provide a
significant source of dietary fiber and micronutrients, along with
a substantial complement of phytochemicals including those
thought to have a role in reducing the risk of chronic diseases
such as atherosclerosis, obesity, diabetes, and cancer. This article
presents a compilation and discussion of the diverse array of
major bioactive compounds that reside in wheat bran. These
compounds in wheat are generally qualitatively similar across
different cereal grains, but specific proportions and distribution
in the kernel can be very different. The health efficacies of these
compounds are not substantially discussed as several excellent
reviews or books on the topic are available for further reading.
Structure of the Wheat Kernel and the Distinctiveness
of Bran
Like all cereal grains, the wheat kernel is defined botanically as a
caryopsis, that is, it contains the seed for generation of a new
plant, plus external tissues, collectively referred to as the pericarp,
which is fused to the seed coat and represents the ripened ovary
http://dx.doi.org/10.1016/B978-0-12-394437-5.00109-1
 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran 269

Hairs of
brush

Endosperm
(~82%)
Crease

Aleurone cells (6-9%)

Nucellar tissue
Seedcoat (testa)
Tube cells Bran
Cross cells Inner and outer (~15%)
Hypodermis pericarp
Epidermis

Germ
(3%) Scutellum

Figure 1 The structure of the wheat kernel sectioned longitudinally (left) and transversely (right). There are six bran layers plus the aleurone, which is
invariably part of conventionally milled bran. The crease, which runs the full length of the kernel, prevents fully efficient separation of bran from
endosperm. Typical weight proportions contributed by bran, germ, and endosperm are shown. Adapted from Dexter JE and Sarkar AK (2004) Wheat: dry
milling. In: Wrigley C, Corke H, and Walker C (eds.), Encyclopedia of Grain Science, pp. 363374. Oxford, UK: Elsevier Science.

Table 1 Proximate and select nutrient composition of the three different components of whole wheat grain, with flour, bran, and germ being
products of commercial millinga

Nutrients Whole wheat (%) Refined flour (endosperm %) Bran (%) Germ (%)
a
Moisture 10.5 11.9 9.9 11.1
Proteina 13.4 10.3 15.6 23.2
Total lipida 2.1 0.98 4.3 9.7
Ash (minerals)a 1.6 0.47 5.8 4.2
Total carbohydrates (by difference)a 72.3 76.3 64.5 51.8
Total dietary fibera 11.5 2.7 47.8 13.2
Sugarsa 0.41 0.27 0.41 37.6
Total bioactive phytochemicalsb 1.2 nd 6.2 nd
a
Data source: USDA National Nutrient Database for Standard Reference (US Department of Agriculture, 2015). Carbohydrates includes total dietary fiber (TDF), starch, and sugars; the
sugar value for germ carbohydrates TDF.
b
Values for whole wheat and bran (not determined, nd for endosperm and germ) correspond to the sum of major phytochemicals considered and reported in this article in descending
order of content in bran: phytic acid, betaine, ferulic acid ferulic acid dehydrodimers, alkylresorcinols, choline, phytosterols, sum of other phenolic acids, total B vitamins, and
lignan.

wall of a wheat floret. The pericarp, which represents about 50%
of the thickness of bran, should not be confused with the hull or
husk of the grain. Modern wheat has evolved as a free-threshing
grain, that is, the caryopsis is naked and does not include the
highly fibrous hull, which is easily removed as chaff in mature
wheat when threshed or harvested. In this way, wheat is distinct
from some other grains like most barley types, as well as oats and
rice, whose kernels naturally include the hull.
The three major components of the wheat kernel are the
germ, bran, and endosperm (Figure 1). Based on research
involving careful wheat kernel dissection, endosperm, bran
(pericarp, testa, and aleurone), and germ (embryo and
scutellum) have been found to represent about 83%,
1315%, and about 3% by weight of the whole grain, respec-
tively. These three fractions of the wheat kernel are relatively
distinct in basic chemical composition (Table 1). The endo-
sperm is energy-rich as it is predominantly composed of starch
but contains less than 10% of the nutritive value (minerals,
vitamins, and dietary fiber) of whole grain. Germ is distinct in
its relatively high protein and lipid content and exceptionally
high concentrations of sugars. Bran contains nearly all the
bioactive compounds of wheat, that is, it is enriched in fiber
components (nonstarch polysaccharides and lignin), minerals,
B vitamins, and other phytochemical compounds (Table 2).
270 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran

From the seeds perspective, the germ contains the embry-
onic plant and an organ called the scutellum, which separates
the embryo from the endosperm. During germination of the
seed, the scutellum absorbs nutrients from the endosperm to
provide food for the germinated seed and growing seedling.
The biological function of bran is very different, which reflects
its complex and multilayered structure as well as chemical
composition; it is to protect the seed from microorganisms
and insects and from adverse weather conditions.
The wheat kernel, as with all cereals, contains a crease that
runs parallel to its long axis, which covers about 25% of its
surface. This crease affects the utilization of wheat for food,
such that the endosperm can only be freed from bran by
conventional roller milling using relatively low flour extraction
rates. The low extraction rate, in the context of milling, mini-
mizes significant contamination of the flour by bran and
germ and results in the production of refined flour, which is
used to make the so-called white bread. Low extraction rates
in flour milling also result in the exclusion of the full comple-
ment of the great many bioactive compounds, which are
Table 2 Average concentration of phytochemicals enhanced in
wheat bran compared to whole grain (mg/100g)a
Phytochemicals
Lignan
Phytosterols
Betaine
Choline
B vitamins, total
B1 (thiamin)
B2 (riboflavin)
B3 (niacin)
B5 (pantothenic acid)
B6 (pyridoxine)
B9 (folate)
a
Values are means of range data in Table 2 of Fardet A (2010) New hypotheses for the
health-protective mechanisms of whole-grain cereals: What is beyond fibre?
Nutrition Research Reviews 23: 65134.
100 Thiamine
Proportion of vitamin retained (%)
concentrated in commercial wheat bran, which includes the
bioactive-rich aleurone layer that represents about 50% by
weight of total bran (Figure 1).
It is important to point out that commercial wheat bran is
not the same as botanical bran, that is, as it exists in the original
grain, which excludes the aleurone layer, which developmen-
tally forms the outer part of endosperm (Figure 1). Wheat bran
can be derived from various stages of the milling process and,
depending on the setup of the mill in terms of flour extraction,
can vary in purity, that is, endosperm contamination, and
hence can vary in its content of fiber, starch, protein, and
other constituents, due to milling effects alone and also due
to genotype, wheat hardness, and environmental conditions
during wheat kernel development. Wheat bran is therefore not
a standardized product of milling and can be produced from
conventional dry roller milling or a combination of conven-
tional milling and abrasion processing similar to that used in
rice polishing. An example of how the degree of flour extrac-
tion in commercial milling affects the retention of the B vita-
min group of bioactive compounds is shown in Figure 2. A
milling extraction rate of 100% is the equivalent of whole
wheat flour.
Nature of Variation in the Content of Bioactive
Whole grain Bran
Components in Bran
0.4 4.75
78 158 It is clear from the literature that there is considerable variation
156 868 in the content of bioactive components in wheat or bran
111 172 depending on the type of wheat or variety (genotype effect)
9.02 33.5 and the environment in which the wheat is grown. Recent
0.56 0.65 science has nicely documented this variation for European
0.18 0.51 wheats and has shown that whole grains can vary from about
6.5 28
1.5- to 4-fold depending on the specific compound and more
1.35 3.15
importantly that some compounds have ample heritability to
0.38 1.0
0.05 0.23 allow breeders to further increase levels if that is desired.
Regarding bran, however, there is another source of variation
that creates issues in accurately documenting bioactive compo-
nent content, that is, milling. As noted in the previous section,
wheat bran is not a standardized product of milling. While it is
possible to obtain pure bran or even isolate specific bran
tissues by hand dissection, this is not the form of bran available

to consumers. It is impossible to obtain 100% pure wheat bran

Pantothenic acid
by any manner of mechanized milling or grinding procedure
Riboflavin
80 due to the tight bond that exists between the aleurone cell layer
Folic acid
Niacin of the wheat kernel and endosperm cells that lie immediately
60 Pyridoxine inward of the aleurone. Depending on the type of wheat or
genotype, and milling procedure, roller-milled bran can con-
40 tain from 5% to 20% of adhering endosperm, which is
enriched in starch. Typically, the harder the wheat, the more
20 pure the resulting bran will be. This variation in endosperm
content of milled bran accounts in part for the wide variation
0 in content of fiber and bioactive components that has been
40 50 60 70 80 90 100 reported in the scientific literature.
Milling extraction rate (%)

Figure 2 The effect of wheat roller milling on the retention of selected B

vitamins in flour. The figure illustrates the invariably uneven
distribution of bioactive compounds in the wheat kernel using B vitamins
as an example. Adapted from Figure 8 of Akyroyd WR and Doughty J
(1970) Wheat in Human Nutrition. Rome: FAO.
Structure of Bran and the Distribution of Bioactive
Components
Bran represents the outermost portion of the grain, which sur-
rounds the starchy endosperm. Bran can be divided into
 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran
nucellar tissue &
seed coat
outer
pericarp aleurone
cross
cells
endosperm
Figure 3 Fluorescence microscopy image showing the outer
endosperm and different tissue layers of bran revealing the distribution of
ferulic acid and other minor phenolic acids in wheat bran and endosperm.
Aleurone contains the highest concentration of phenolic acids in the
wheat kernel. Bar 50 mm. Adapted from Ndolo VU, Beta T, and Fulcher
RG (2013) Ferulic acid fluorescence intensity profiles and concentration
measured by HPLC in pigmented and non-pigmented cereals. Food
Research International 52: 109118, with permission.
different tissue layers, more than what is depicted in Figure 1.
Accordingly, bran is the most complex physical component of
the wheat kernel. Bran basically comprises the outer and inner
pericarp, seed coat (often referred to as testa), and a layer of large
and thick-walled cells called the aleurone (Figures 3 and 4). In
the mature wheat seed, the aleurone consists of living tissue, one
cell thick that surrounds the endosperm. During germination,
aleurone cells have an important enzyme secretion function;
they respond to the embryos release of the hormone gibberellic
acid, to synthesize and secrete enzymes such as a-amylase and
proteases, which hydrolyze endosperm starch and protein to
sugars and amino acids, respectively, to support the early growth
of the embryo into a developing seedling.
Quantitatively, the predominant bioactive component of
wheat bran is fiber. Fiber is not a distinct chemical compound,
but a heterogeneous mixture of various nonstarch polysaccha-
rides plus lignin and other compounds. Ranked in order of
decreasing content, the major bran fiber bioactive compounds
are arabinoxylans, cellulose, lignin, fructans, and b-glucans
(Figure 5). Together, these compounds comprise over 90% of
the fiber content of wheat (Figure 5) and from 45 to 55% of
the fiber content of bran. In contrast, refined wheat flour
depending on type (e.g., cake flour or all purpose) contains
from 1.5 to 3% fiber (Table 1).
Lignin is molecularly different from the other fiber compo-
nents as it is a complex polyphenolic compound that does not
contain any sugar molecules. All the fiber components of bran
reside in the cell walls of the tissue layers (e.g., Figure 4, which
depicts the distribution of arabinoxylans and b-glucans in
aleurone cell walls) and include small molecules such as phe-
nolic acids (Figures 3 and 6). Wheat bran phenolic acids and
271
Figure 4 Double labeling of arabinoxylan (AX) and b-glucans in
aleurone cell walls from wheat grain. Polyclonal antixylan and
monoclonal anti-b-glucan antibodies were used with a second-stage goat
antirabbit (orange-red fluorescence) and a second-stage goat antimouse
(green fluorescence) antibody, respectively. Yellow fluorescence
indicates the presence of both b-glucans and AX. The light-colored
circular structures within the vacuoles of the aleurone cells are storage
protein bodies that contain dark-stained inclusion bodies or globoids in
which the phytochemical phytin is concentrated. Reprinted from Saulnier
et al. (2007) Plant cell wall polysaccharides in storage organs: Xylans
(food applications). In: Kamerling J, Boons G-J, Lee Y, Suzuki A,
Taniguchi N, and Voragen AGJ (eds.) Comprehensive Glycoscience, vol.
2, pp. 653689. Oxford: Elsevier Science Ltd.
30
24.0
25 Whole wheat
21.4
20 Bran
g/100 g
15
10
5.9 5.2
5 3.9
2.0 2.3 2.3 2.3 2.0
0
Figure 5 Components of fiber in whole wheat and bran. Data
compiled from a variety of sources showing typical values averaged
across wheat genotypes and/or samples.
ferulic acid most notably are important bioactive phytochem-
icals that have a structural role to cross-link the nonstarch
polysaccharides with other cell wall components including
lignin, thereby increasing the physical integrity of cell walls,
and also provide a barrier to invading pathogenic fungi and
other microorganisms, which are naturally present in the peri-
carp of wheat bran. Similarly, the seed coat that separates the
pericarp from aleurone and endosperm appears to be chemi-
cally inert and waterproof. This latter property of the seed coat
272 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran

500 25
410 Whole wheat
400 20.0 Whole wheat
Bran 20
296 Bran

mg/100 g
mg/100 g
300 15
200 10 9.0
117 6.3
73 5 3.7 3.5 3.2
100 2.2
1.5 1.3 0.7
0 0
Ferulic + dimers Alkylresorcinols
(a)

(b)

Figure 6 Phenolic compounds significantly enhanced in wheat bran compared to whole grain. Ferulic acid plus ferulic acid dehydrodimers and
alkylresorcinols (a) are the predominant phenolic compounds, each having at least tenfold higher concentration compared to the total of all other
phenolic compounds including simple phenolic acids (b) and flavonoids (not shown). Phenolic acid data compiled from a variety of sources.
Alkyresocinol values are the mean of those reported by Fardet (2010). Reproduced from Mattila et al. (2005) Contents of phenolic acids, alkyl- and
alkenylresorcinols, and avenanthramides in commercial grain products. Journal of Agricultural and Food Chemistry 53: 82908295.

likely derives at least in part from the presence of a distinct

group of phenolic lipid compounds called alkylresorcinols,
which contain long hydrophobic alkyl chains similar to satu-
rated fatty acids.
Botanically, the aleurone layer of wheat is part of the endo-
sperm, but because of its high adherence to the seed coat,
which is likewise strongly bonded to the pericarp, aleurone is
mainly found in bran fractions after milling. Wheat as noted
has a single-cell aleurone layer. In contrast, barley aleurone has
two to four cell layers. The thicknesses of wheat brans tissue
layers also vary. The aleurone is the thickest layer (up to
65 mm), a feature that can be clearly seen in microscope images
(e.g., Figures 3 and 4). The outer pericarp is of intermediate
thickness (1530 mm), and the seed coat is the thinnest layer
(58 mm). Variation in bran thickness in an individual wheat
grain is believed to be due to the irregularities of thickness of
aleurone cells affecting the separation of bran from endosperm
during milling. Thus, flour yield during milling depends in
part on bran thickness, size of grain (bran and endosperm
content), and crease characteristics. An interesting question is
whether reported differences among different wheats in phe-
nolic acid content and antioxidant capacity are due, at least in
part, to variation in thickness of the aleurone layer and there-
fore concentration of constituent phytochemicals such as feru-
lic acid. There appears to be no science to answer this question
at the present time.
From a bioactive phytochemical perspective, the aleurone is
the most significant tissue in the wheat kernel. It contains in its
thick cell walls relatively high levels of fiber components
(Figure 4). Aleurone cell walls are also rich in phenolic com-
pounds such as ferulic acid and r-coumaric acid (Figure 3) and
therefore contribute the highest antioxidant capacity of any
wheat tissue. Aleurone cells are also enriched in B vitamins
(B3, or niacin; B2, or riboflavin; B5, or pantothenic acid; B6, or
pyridoxine; and B9, or folate), minerals (e.g., phosphorus,
potassium, magnesium, manganese, and iron), and choline
and betaine to a lesser extent. In addition, wheat aleurone
has a very high content of phytic acid, which is the
5
Whole wheat 4.2
4 Bran
g/100 g
3
2
1 0.7
0
Phytic acid
Figure 7 Phytic acid content in whole wheat and bran. Values are the
mean of those reported by Fardet (2010).
predominant storage form of phosphorus in the seed. Phytic
acid, also known as inositol hexaphosphate, is the phytochem-
ical in highest concentration in whole wheat (0.31.5%) and
wheat bran (2.36%) (Figure 7). Phytic acid is typically
complexed in the seed with minerals such as calcium or mag-
nesium, and in this form, it is called phytin. Phytin exists
within the cavity or vacuole of aleurone cells and can be seen
in microscopic images (e.g., Figure 4) as numerous dark-
stained inclusions within circular storage protein bodies.
While phytate has been traditionally viewed as an antinutrient
because of its mineral-binding properties, it is also known to
have compelling anticancer properties.
Molecular structures of several of the major bioactive com-
pounds in bran as discussed are shown in Figure 8.
Brief Descriptions of Some of the Major Bioactive
Compounds in Wheat Bran
A common physiological role for many of the fiber-bound
bioactive compounds that are enhanced in wheat bran, nota-
bly nonstarch polysaccharides, oligosaccharides (fructans),
 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran 273

OH
-O O
OH P P O-
O
O O
O
P O-
O
R O O- O-
P
O O-
R = CnH2n+1 O O
n = 15, 17, 19, 21, 23, 25 O P O-
OH O P O-
O- OH

(a) Alkylresorcinol (b) Phytic acid

OH
OH HO
HO
A
A HO
HO
OH O
O O O O
HO O
O X O X X O X
HO O O O
O OH O O OH
HO A

O
OH
H3CO
O
F Arabinoxylan
(c) HO

R3
Cinnamic acid derivatives
ferulic acid, R1 = OCH3; R2 = OH
R2 CH = CH- COOH sinapic acid, R = OCH ; R = OH
1 3 2
p-coumaric acid, R2 = OH

R1
Phenolic acids
R3

Benzoic acid derivatives

R2 COOH vanillic acid, R1 = OCH3; R2 = OH
syringic acid, R1 = R5 = OCH3; R2 = OH
p-hydroxybenzoic acid, R2 = OH
R1
(d)

Figure 8 Basic molecular structures or portions of the structure of four types of bioactive compounds in wheat bran: alkylresorcinol (a); phytic acid (b);
arabinoxylan (c) showing xylose (X) backbone with substituted arabinose (A) molecules, some of which are linked to the phenolic acid ferulic acid (F); phenolic
acids (d). (Figure 8c adapted from Saulnier et al. (2007) Plant cell wall polysaccharides in storage organs: Xylans (food applications). In: Kamerling J,
Boons G-J, Lee Y, Suzuki A, Taniguchi N, and Voragen AGJ (eds.) Comprehensive Glycoscience, vol. 2, pp. 653689. Oxford: Elsevier Science Ltd.)

and lignin (Figure 5), is to promote intestinal health. There are
an amazing number of bacterial cells in the colon in particular,
and estimates run from 1011 to 1012 cells per gram. This huge
population of cells, often referred to as the microbiome, is
gaining considerable importance in science due to an increased
understanding of the profound health effects, both positive
and negative, that good or poor gut health, respectively, can
contribute.
Wheat bran (fiber) is particularly relevant in this discussion
as it is highly indigestible, yet fermentable by bacteria in the
colon, which have enzymes necessary to hydrolyze the poly-
meric compounds in bran such as polysaccharides. It has been
known for a long time that the stimulation of fermentation
caused by consuming these fiber bioactives leads to an increase
in bacterial mass and consequently fecal mass, which has a
positive effect on laxation and regularity. What is only being
274 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran
understood very recently is that fermentation causes alterations
in the microbiome, which in turn alters bacterial metabolites
(short-chain fatty acids), modulates inflammatory responses,
and likely leads to a decreased risk of inflammation-associated
and other metabolic abnormalities such as atherosclerosis and
type II diabetes. Also, by producing short-chain fatty acids,
most notably butyric acid, and other metabolites, the intestinal
microbiome is believed to contribute to a gut environment that
resists the proliferation of pathogenic bacteria.
Arabinoxylans and Ferulic Acid
Arabinoxylans, which are part of a group of nonstarch poly-
saccharides compounds, originate in the cell walls of bran
tissues. Pentosans, a term still used in the current literature
synonymous with arabinoxylans, represent nonstarch poly-
saccharides composed of five-carbon or pentose monosaccha-
rides, xylose, and arabinose (Figure 8). More precisely,
arabinoxylans consist of a xylose polymer backbone with
frequent branching of arabinose residues. Significantly from
the standpoint of structure and bioactivity, arabinoxylans
contain the phenolic compound ferulic acid, which is cova-
lently bound to arabinose residues. Arabinoxylans are quan-
titatively the most important bioactive compound in wheat
and can represent as much as 30% of the content of bran
(Figure 5).
Ferulic acid is the main cell wall-bound phenolic acid in all
cereal grains and in wheat it is found almost exclusively asso-
ciated with arabinoxylans. Arabinoxylans are very diverse in
structure and relative content of arabinose, xylose, and ferulic
acid molecules. They differ in this way from those found in the
endosperm, which tend to be more soluble than arabinoxylans
in bran, which are highly insoluble. Arabinoxylans are known
to have a prebiotic function in the gut, that is, they can stim-
ulate the growth of healthful bacteria such as Bifidobacterium
and, in this way and others, contribute to gut health. The so-
called dehydroferulic acid dimers (diferulates) have an impor-
tant structural role to cross-link and therefore stabilize arabi-
noxylan polysaccharides and hence the cell walls in bran.
Consequently, diferulates may be partly responsible for the
insoluble nature attributed to the dietary fiber of wheat bran.
Importantly, the antioxidant potency of extracts of wheat and
bran is predominantly determined by the high content of
ferulic acid in bran (Figure 6), which is enriched in the aleu-
rone fraction. The antioxidant capacity of biomolecules such as
ferulic acid is associated with the scavenging of free radicals,
many of which are reactive oxygen molecules, which are often
toxic to cells causing inflammatory responses and damage to
lipids, proteins, and nucleic acids.
b-Glucan
b-Glucans are so-called b-linked polysaccharides of glucose
monomers. b-Glucans are a diverse group of molecules that
can vary with respect to molecular mass, solubility, and viscos-
ity. In wheat, b-glucan is found mainly in the aleurone cell
walls of wheat bran but are distributed more widely in the
grain of other cereals. In oats, b-glucans are concentrated in
the subaleurone layer, while in barley and rye, b-glucans are
located mainly in the endosperm. The b-glucan levels of wheat
bran are low (2%) and represent <1% of the total dietary
fiber content of whole wheat. Because of its low concentration
in wheat, b-glucans likely contribute relatively little by way of
health attributes, compared to oats and barley. In these latter
grains, b-glucan is considered to be an important soluble fiber
that is associated with lowering serum cholesterol levels and
attenuating glycemic response. These effects originate in the
upper gastrointestinal tract and probably are related to the
viscosity of b-glucan.
Cellulose
Cellulose is a very abundant cell wall polysaccharide in the
outer pericarp and intermediate layers of wheat bran. Cellulose
consists of glucose units linked together by b-14 glycosidic
bonds to form a linear polysaccharide. Cellulose comprises
about 21% of wheat bran on a 14% moisture basis
(Figure 5), which is substantially enhanced compared to
whole wheat at (2%); however, estimates vary widely. Cellu-
lose is considered to be very resistant to intestinal microbial
fermentation when compared with the noncellulosic fiber
polysaccharides such as arabinoxylans and especially b-glucans
whose digestibility is relatively high. Nevertheless, bacteria in
the colon are known to contain cellulase enzymes, which can
break down cellulose leading to its fermentation and likely
beneficial effects for health.
Lignin
Lignin is a complex polyphenolic polymer residing in the
pericarp and seed coat cell walls of wheat bran. Lignin is a
nonpolysaccharide cell wall component that is a polar poly-
mer. Whole wheat has a lignin concentration of 2% of dry
material that is only moderately enhanced in wheat bran to
levels of about 5% (Figure 5). The distribution of lignin in
wheat has been reported to range from about 2 to 9% in
different bran layers but is not found in aleurone according
to the literature. Ferulic acid, together with dehydroferulic acid,
is a component of lignocelluloses, serving to cross-link the
lignin and polysaccharide, thereby conferring rigidity to the
cell walls. Like cellulose, lignin is fermented at low levels by
bacteria in the colon and is able to increase fecal bulk. Also,
lignins lower intestinal transit time, which is important to
reduce exposure time of dietary carcinogens adsorbed in
fiber. Lignins also possess in vitro antioxidant activity and
may mitigate oxidative DNA damage and development of
diseases in relation to increased oxidative stress.
Fructans
Fructans are a category of functional food carbohydrates that
encompass naturally occurring plant oligosaccharides and
polysaccharides containing fructose. Fructans are different in
 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran 275

nature and classified according to linkage and origin. Fructoo- Compared to simple phenolic acids, alkylresorcinols represent
ligosaccharide is a subgroup of inulin, consisting of linear a very distinct group of phenolic compounds in terms of molec-
chains of fructose sugar monomers linked by b-(12) bonds ular properties, that is, they are amphiphilic, meaning they have
with a low degree of polymerization (10). Whole wheat grain both polar and nonpolar properties. This bioactive compound
and bran fructan contents are similar to those reported for consists of a phenolic ring with two hydroxyl groups and a
lignin. Fructans are classified as a functional fiber that are hydrophobic and mostly saturated alkyl chain that can contain
believed to be extensively fermented, but with little effect on from 15 to 25 carbons (Figure 8). Wheat bran is a rich source of
stool weight. Fructans are known to have positive effects on alkylresorcinols and has been reported to contain levels of about
calcium uptake in the gut and hence calcium bioavailability. 0.3%, which is fourfold higher than its content in whole wheat
(Figure 6). Refined flour contains essentially no alkylresorcinol
content. Accordingly, alkylresorcinols can be used as biochem-
Phenolic Acids
ical markers of whole wheat and bran in foods.
Several studies have reported that alkylresorcinols have the
Phenolic acids are a very noteworthy constituent of wheat
ability to protect cellular lipid components from oxidative pro-
bran. Apart from flavonoids, which are found to a large extent
cesses. Accordingly, alkylresorcinols appear to behave as antiox-
in wheat germ and in relatively low concentration in bran,
idant compounds, but they are generally known as weak
virtually all other phenolic compounds (phenolic acids and
antioxidants, based on their hydrogen-donating and radical-
phenolic acid esters) reside in wheat bran tissues, most notably
scavenging abilities, compared to some other bioactive com-
aleurone. Phenolic acids can be subdivided into two major
pounds such as a-tocopherol (vitamin E). Their antioxidant
chemical groupings: hydroxybenzoic acids and hydroxycin-
potential depends on their chain length, which affects their
namic acids (Figure 8). Hydroxybenzoic acids of significance
amphiphilic behavior and incorporation into cell membranes;
in bran, although levels are relatively low (Figure 5), include
studies have shown that the shorter the chain length, the higher
vanillic acid, synergic acid, and r-hydroxybenzoic acid. They
the inhibitory level of lipid oxidation. Also, membrane incorpo-
are typically components of complex compounds like lignins
ration of alkylresorcinols could accommodate a biological effect
and tannins. The concentration of hydroxybenzoic acids in
given their high concentration in wheat and bran, but limited
wheat and bran is low compared to hydroxycinnamic acids,
data on alkylresorcinols effective dose levels require further
which includes r-coumaric, and ferulic acid, which form links
studies on risk modulation of cancer and cardiovascular disease.
to cell wall structural components such as arabinoxylans,
Alkylresorcinols are absorbed through the lymphatic system and
cellulose, and lignins through ester bonds.
have been recovered from rat adipose tissue, human lipoproteins
By far, the predominant phenolic acid in wheat is (trans)
(the main transporter), human erythrocyte membranes, and
ferulic acid, which comprises more than 90% of total phenolic
human plasma. Animal studies have shown that alkylresorcinol
acids in wheat bran. Wheat bran is one of the richest sources of
is highly bioavailable, which indicates that apparently, the
ferulic acid with levels as high as 1400 mg/100g being
indigestible bran matrix does not greatly inhibit absorption.
reported. Ferulic acid is predominant in wheat aleurone,
pericarp, and embryo cells, while only trace amounts can be
found in starchy endosperm. Aleurone cells are associated with
the highest concentration of ferulic acid (Figure 3). The seed
Phytic Acid
coat layer of wheat bran contains significant quantities of
ferulic acid, higher than those in the pericarp. Total ferulic
Perhaps the most notable phytochemical constituent of wheat
acid concentration in wheat bran is typically about four times
bran is phytic acid, which is composed of a simple sugar (inosi-
higher than that in whole wheat grain (Figure 5).
tol) with phosphate groups attached to each of the six carbons in
There is considerable scientific evidence that ferulic acid, like
the sugar molecule (Figure 8). It is typically found in food sources
other phenolic compounds, has antioxidant, anticarcinogenic,
high in fiber content, although wheat bran may be the richest
and anti-inflammatory properties. The great majority of these
source in nature. Phytic acid represents the main storage form of
studies are based on animal or cell models. The types of antiox-
phosphorus in wheat and other cereal grains. Phytic acid repre-
idant activities possessed by phenolic acids include radical scav-
sents the most highly concentrated phytochemical in wheat and
enging, metal ion chelation, and inhibition of prooxidant
wheat bran in particular with average levels of 0.7% and 4%,
enzymes. Ferulic acid has been extensively studied for its
respectively (Figure 7). By contrast, endosperm tissue of wheat
in vitro antioxidant effects and has been shown to be effective.
contains very little phytic acid. Phytic acid has traditionally been
However, in vivo studies of phenolic acids from cereals like
considered to be an antinutrient associated with its ability to bind
wheat and fractions like wheat bran are few. Cereal grains such
minerals with positively charged ions (i.e., cations) such as iron,
as wheat that have significant quantities of ferulic acid and other
as well as magnesium and calcium. This binding of minerals
phenolics require in vivo studies to determine if high enough
therefore reduces their bioavailability, but only for poorly nour-
levels of these bioactive compounds become bioavailable when
ished people consuming predominantly whole grains in their
consumed in food to provide health benefits.
diet. Many animal studies have shown phytic acid to be capable
of antineoplastic activities on multiple types of cancers of the
Alkylresorcinols breast, colon, liver, prostate, skin, and skeletal muscle. Potential
mechanisms suggested for phytic acids anticarcinogenic proper-
Alkyresorcinols are phenolic lipids that are present exclusively in ties include cell cycle inhibition, antioxidant ability (metal chela-
bran and specifically in the outer layer of the seed coat. tion), gene alteration, and increased natural killer cell activity.
276 BIOACTIVES AND TOXINS | Bioactive Compounds in Wheat Bran
Summary and Conclusions
The nature of wheats value for human health derives from its
structure, biochemical composition, and functionality of its
components. Because wheat consumption as bread and other
foods is so significant, whole wheat and the bran it contains
have the potential to deliver to the diet considerable amounts
of fiber and constituent bioactive compounds whose function-
ality extends well beyond the role of nutrients, which are
essential for growth and maintenance of life. There is now
ample scientific evidence that consumption of adequate
amounts of bran in whole wheat is associated with lowering
the risk of many chronic diseases, although the specific actions
of individual biologically active compounds in bran have been
only studied using animal and cell models. A more thorough
discussion of the potential health efficacies of the major bio-
active compounds of wheat bran is well beyond the scope of
this article. Many excellent books or review papers have been
published and a few are offered at the end of this article as
suggestions for further reading. These articles are based on a
foundation of hundreds of research studies, and the science in
this area continues to expand in scope and knowledge. Most
exciting perhaps is the microbiome-related research on the
fermentation of fiber and bran that is gaining momentum.
Wheat bran is generally considered to be one of the best, if
not the best, sources of fiber for fecal bulking and laxation. This
sort of bioactivity represents the traditional health benefit
ascribed to bran, but it also suggests that wheat bran interacts
strongly with the microbiome and that this area of research has
enormous potential for greater understanding of mechanisms
underlying the more compelling health efficacies generally
associated with the consumption of whole wheat and bran
and its constituent bioactive compounds.
Exercises for Revision
Are there wheat milling procedures that can generate bran
of higher purity that is more concentrated in bioactive
compounds?
What is the function of the bioactive compounds in wheat
bran from the perspective of the wheat seed?
How much wheat bran is typically produced in a large
milling operation and how much of that bran is destined
for food use and how much for feed? How do these pro-
portions vary in major wheat producing countries?
Are there wheat genotypes or varieties that have bran with
enhanced levels of bioactive compounds?
How does the wheat growing environment affect the con-
tent of bioactive compounds in bran?
Exercises for Readers to Explore the Topic Further
How do bioactive compounds in the bran or whole grain of
other cereals compare to wheat, especially oats, given its
reputation for having dietary benefits?
Select a few fruits or vegetables that are commonly associ-
ated with health benefits. How similar or different are the
bioactives in those foods compared to wheat bran?
Apart from eating whole wheat bread or pasta, think of
other ways to increase your consumption of wheat bran
that would appeal to you.
See also: Bioactives and Toxins: Bioactives: Antioxidants;
Carbohydrates: Cereals: Chemistry and Physicochemistry of Non-
starchy Polysaccharides; Food Grains and the Consumer: Grains
and Health; Food Grains and Well-being: Functional Foods:
Dietary Fibers, Prebiotics, Probiotics, and Synbiotics; Functional
Foods: Overview; Grain Composition and Analysis: The
Composition of Food Grains and Grain-Based Products; The
Basics: Grain: Morphology of Internal Structure.
Further Reading
Anson NM, Hemery YM, Bast A, and Haenen GR (2012) Optimizing the bioactive
potential of wheat bran by processing. Food and Function 3: 362375.
Fardet A (2010) New hypotheses for the health-protective mechanisms of
whole-grain cereals: What is beyond fibre? Nutrition Research Reviews
23: 65134.
Marquart L, Jacobs Jr. DR Jr., McIntosh GH, Poutanen K, and Reicks M (eds.) (2007)
Whole Grains and Health. Oxford: Blackwell.
Okarter N and Liu R (2010) Health benefits of whole grain phytochemicals. Critical
Reviews in Food Science and Nutrition 50: 193208.
Shewry PR, Charmet G, Branlard G, et al. (2012) Developing new types of
wheat with enhanced health benefits. Trends in Food Science and Technology
25: 7077.
Slavin J (2013) Fiber and prebiotics: Mechanisms and health benefits. Nutrients
5: 14171435.
 Bioactives: Antioxidants
T Beta, University of Manitoba, Winnipeg, Manitoba, Canada
KG Duodu, University of Pretoria, Pretoria, South Africa
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Grain bioactives that exert antioxidant effects include phe-
nolic compounds, carotenoids, tocols, sulfur compounds,
betaine, choline, policosanol, melatonin, g-oryzanol, sapo-
nins, and phytates.
Chemical and biological measurements of antioxidant
properties demonstrate varying degrees of activity among
grains, grain fractions, and constituents.
Grain bioactive antioxidants are proposed to potentially
ameliorate the fundamental processes of oxidative stress
that tend to be precursors for many chronic diseases of
lifestyle.
The mechanisms of action of grain bioactives are proposed
to be mainly via either exertion of antioxidant effects or
modulation of the activity of various enzymes involved in
the cellular and molecular pathways and reactions associ-
ated with chronic diseases.
Learning Objectives
To achieve understanding of the major bioactive com-
pounds in grains that exert antioxidant effects when mea-
sured using chemical and biological assays.
To achieve an understanding of the role of grain bioactive
compounds in offering antioxidant protection against
major chronic diseases and the mechanisms of action of
the bioactives.
Introduction
In addition to major and minor nutritional components, cereal
grains including maize, rice, wheat, barley, sorghum, rye, oats,
triticale, buckwheat, and millets contain a wide range of bio-
active phytochemicals associated with antioxidant effects.
Grain phenolics, carotenoids, tocopherols, phytates, and phy-
tosterols are among the list of candidates displaying antioxi-
dant properties. Some bioactive antioxidants are unique to
certain grains such as avenanthramides in oats; xanthophylls
in yellow endosperm maize, wheat, and, sorghum; and antho-
cyanins in blue, red, and purple grains. The literature on anti-
oxidant effects of grain bioactives is not easily comparable due
to the lack of uniformity of the assays in use. However, there
has been an attempt to establish systematic databases such as
the Phenol-Explorer database on polyphenol content of foods.
In vitro antioxidant measurements have been applied to a lesser
extent on non-phenolic bioactive constituents in grains, while
in vivo measurements using humans are largely lacking. Since
grains are staples around the world, they are a notable source
Encyclopedia of Food Grains, Second Edition http://dx.doi.org/10.1016/B978-0-12-394437-5.00110-8
of bioactive antioxidants albeit in relatively lower levels com-
pared to other food sources.
Cereal Grains and Grain Products
Cereal grains include maize, rice, wheat, barley, sorghum, rye,
oats, triticale, buckwheat, and millets. The glumes/husks cover
some grains such as oats. Once dehusked, the cereal caryopsis
is made up of the endosperm, which includes the aleurone
layer, germ, testa or seed coat, and outer pericarp layers as the
main botanical parts. Carbohydrates, proteins, and lipids make
up the major chemical components while minerals and vita-
mins are considered minor nutrients in grains. Phytochemicals
including bioactives in grains make up the remainder of the
minor constituents. The distribution of nutrient components
and phytochemicals varies among the different botanical parts
of the grain as exemplified by whole-grain wheat, wheat bran,
and wheat germ (Table 1).
Following harvest, cereal grains are stored and processed for
human food. Grain processing includes dry and wet milling,
boiling, lime cooking, steaming, baking, frying, extrusion, flak-
ing, roasting, fermentation, malting or sprouting, and brewing.
Storage and processing affect the chemical constituents of
grains including bioactives resulting in an alteration of the
phytochemical profiles of the resultant products. Qiu et al.,
for example, demonstrated a much lower total phenolic con-
tent in quick-cooking wild rice than in uncooked wild rice
varieties. Whole-grain cereal, germ, and bran (aleurone, testa,
and outer pericarp layers) obtained during dry milling or hand
dissection often display higher phytochemical levels than the
endosperm. Several phytochemicals, exerting antioxidant
effects, have been identified in grains. Consumption of
whole-grain cereals can potentially protect the human body
against increased oxidative stress.
Reduction and/or Prevention of Oxidative Stress
Cellular redox processes in the body lead to the formation of
highly reactive free radicals, generally in the form of reactive
oxygen species or reactive nitrogen species. Some examples of
such radicals that are of importance in living organisms
include hydroxyl (OH ), superoxide (O2
), nitric oxide
l l
l l
(NO ), and peroxyl (RO2 ). It is postulated that at high con-
centrations, these reactive species cause the condition known
as oxidative stress, which is harmful to cell structures. Oxida-
tive stress is a term used to describe the condition where there
is an imbalance between the generation of reactive oxygen or
nitrogen species and the activity of the natural antioxidant
defense systems of the body. Severe oxidative stress can cause
cell damage and death, and it is implicated in various human
277
278 BIOACTIVES AND TOXINS | Bioactives: Antioxidants

Table 1 Antioxidants in whole-grain wheat, wheat bran, and wheat germ

Bioactive: antioxidant Whole-grain wheat Wheat bran Wheat germ

Reduced glutathione 1.045.74 mg/100 g 1.719.4 mg/100 g 19.4245.7 mg/100 g

Methionine 0.170.24 g/100 g 0.200.29 g/100 g 0.390.58 g/100 g
Cystine 0.190.40 g/100 g 0.320.45 g/100 g 0.350.61 g/100 g
Lignins 0.92.8 g/100 g 2.29.0 g/100 g 1.31.6 g/100 g
Phytic acid 0.281.50 g/100 g 2.36.0 g/100 g 1.32.2 g/100 g
Fe 1.014.2 mg/100 g 2.519.0 mg/100 g 3.910.3 mg/100 g
Zn 0.88.9 mg/100 g 2.514.1 mg/100 g 1018 mg/100 g
Mn 0.97.8 mg/100 g 414 mg/100 g 918 mg/100 g
Se 0.00033.000 mg/100 g 278 mg/100 g 179 mg/100 g
Cu 0.091.21 mg/100 g 0.842.20 mg/100 g 0.701.42 mg/100 g
Mg 17191 mg/100 g 390640 mg/100 g 200340 mg/100 g
Thiamin (B1) 0.130.99 mg/100 g 0.5060.800 mg/100 g 0.82.7 mg/100 g
Tocols (E) 2.37.1 mg/100 g 9.5 mg/100 g 23.131 mg/100 g
Total tocopherols 1.062.89 mg/100 g 2.4 mg/100 g 21.530.6 mg/100 g
Total tocotrienols 1.094.49 mg/100 g 7.1 mg/100 g 1.31.6 mg/100 g
Total carotenoids 0.0440.626 mg/100 g 0.251.18 mg/100 g
b-Carotene 0.0050.025 mg/100 g 0.030.010 mg/100 g 0.062 mg/100 g
Lutein 0.0260.383 mg/100 g 0.0500.180 mg/100 g
Zeaxanthin 0.0090.039 mg/100 g 0.0250.219 mg/100 g
b-Cryptoxanthin 1.1213.28 mg/100 g 0.0180.064 mg/100 g
Total phenolic acids 16102 mg/100 g 7611384 mg/100 g
Bound phenolic acids 1478 mg/100 g 148340 mg/100 g
Total ferulic acid 16213 mg/100 g 138631 mg/100 g 7124 mg/100 g
Total dehydrodiferulic acid 1.576.0 mg/100 g 13230 mg/100 g 9 mg/100 g
Total dehydrotrimer FA 2.63.5 mg/100 g 1525 mg/100 g
Total flavonoids 3043 mg CE/100 g 14.940.6 mg/100 g 300 mg RE/100 g
Anthocyanins 0.4552.60 mg CE/100 g 0.948.0 mg/100 g
Lignans 0.1990.619 mg/100 g 2.86.7 mg/100 g
Alkylresorcinols 11.6128.8 mg/100 g 215323 mg/100 g
Betaine 22291 mg/100 g 2301506 mg/100 g 3061395 mg/100 g
Total choline 27195 mg/100 g 74270 mg/100 g 152330 mg/100 g
Phytosterols 5798 mg/100 g 121195 mg/100 g 410450 mg/100 g
Melatonin 0.20.4 mg/100 g

Source: Fardet A (2010) New hypothesis for the health-protective mechanisms of whole-grain cereals: What is beyond fibre? Nutrition Research Reviews 23: 65134.

diseases including cancer, cardiovascular disease, type 2
diabetes, and even cataractogenesis.
It is hypothesized that bioactives from cereal grains could
offer potential health benefits by protecting against diseases
associated with oxidative stress. A lot of in vitro evidence has
emerged that shows the ability of cereal bioactives to protect
cell constituents against oxidative stress and oxidative damage
of biological tissues. This observed protective effect has been
attributed to various mechanisms. These include the suppres-
sion of the formation of reactive oxygen species by either
inhibiting enzymes or chelating trace elements involved in
free radical production, scavenging reactive oxygen species, or
upregulating or protecting antioxidant defense systems.
Antioxidants from Whole-Grain Cereals
Whole-grain cereals serve as sources of antioxidants that
include sulfur compounds, minerals, B vitamins, phytic acid,
tocols, carotenoids, phenolic compounds, lignins, lignans,
alkylresorcinols, betaine, choline, policosanol, melatonin,
g-oryzanol in rice bran, avenanthramides, and saponins in
oats as reviewed by Fardet. These antioxidants differ in their
structure and mechanism of action, which can be direct or
indirect. Mechanisms may involve direct free radical scaveng-
ing or Fe chelation. Indirect antioxidants such as Fe, Zn, Cu,
and Se act as cofactors of antioxidant enzymes. Very few studies
have examined the antioxidant hypothesis; hence, in vivo inves-
tigations are needed to determine the complex and synergistic
mechanisms taking place in human subjects.
Sulfur Compounds
Total methionine and cystine contents of whole-grain wheat,
wheat bran, and wheat germ are 0.5%, 0.6%, and 1.0%, respec-
tively (Table 1). Other cereal varieties may contain higher levels.
Methionine and cystine contents of commercially prepared
whole-wheat bread based on USDA database are, respectively,
155 and 214 mg/100 g, contributing an average 47 and 64 mg
day
1 when a 30 g slice of bread is consumed. Significant
amounts of at least 280 mg methionine and 380 mg cystine
per day can be obtained by following USDA food guide pyramid
that recommends between 6 and 11 daily servings of whole-
grain cereal products. The two sulfur-containing amino acids are
both precursors of reduced glutathione (GSH), an intracellular
antioxidant that controls cell oxidative status by participating in

gene expression through the modification of the thiol redox
status. GSH is also present at levels of 1.045.74, 1.719.4,
and 19.4245.7 mg/100 g in whole-grain wheat, wheat bran,
and wheat germ, respectively, although its absorption in
humans is deemed negligible in its free form.
Minerals
The contents of Fe, Zn, Mn, Mg, Cu, and Se in whole-grain
wheat, wheat bran, and wheat germ are shown in Table 1 as
reviewed by Fardet. Low levels of 3.8% Fe are absorbed in
humans following the consumption of wheat bran rolls.
About 1720% and 1995% of zinc is absorbed in humans
and rats, respectively, following the consumption of whole-
grain wheat. There is limited literature on apparent absorption
of Mn in humans, although levels are known to reach
1418 mg/100 g in wheat bran and wheat germ compared to
0.0780.079 mg/100 g for selenium in respective fractions.
Studies have shown a 6085% absorption of selenium in rats
following the consumption of wheat and sodium selenite.
Fe, Zn, Cu, and Se act as cofactors of antioxidant enzymes
and therefore serve as indirect antioxidants. Cu exerts its anti-
oxidant effect mainly as superoxide dismutase cofactor. Mg
displays antioxidant effects by acting against lipid peroxida-
tion. Se protects cell membranes from damage caused by lipid
oxidation.
Thiamin and Riboflavin
Both thiamine (B1) and riboflavin (B2) are present in grains
including wheat (Table 1). The two B vitamins exert antioxi-
dant effects in addition to their several well-documented phys-
iological functions.
Phytic Acid
Phytic acid (myoinositol hexakisphosphate) can reach up to 6%
in the bran of certain wheat varieties. It displays strong in vitro
antioxidant activity; however, it has been considered an antinu-
trient due to its ability to chelate minerals such as Zn, Fe, Ca, and
Mg, reducing their bioavailability. Processing procedures such as
the preparation of sourdough to produce bread can activate
endogenous phytases and lower pH, thereby reducing the che-
lation of minerals by phytic acid. Phytic acid can also chelate Fe
and limit the oxidative damage caused by the Fenton reaction,
which would otherwise lead to the production of the very
l
reactive free radical OH . Strategies are required that pre-
hydrolyze phytic acid so as to maximize mineral bioavailability
while simultaneously enhancing its antioxidant effect against
free radicals in the gastrointestinal tract (GIT).
Phytic acid is poorly absorbed in humans. In the absence of
the enzyme phytase, phytate is not likely to be absorbed at the
intestinal level in humans. About 5860% of phytate is
degraded into lower myoinositol phosphates in ileostomates
following the consumption of raw wheat bran and only 5%
following the consumption of phytase-deactivated wheat bran.
As an antioxidant, phytic acid chelates various metals includ-
ing the suppression of damaging Fe-catalyzed redox reactions,
inhibits xanthine oxidase, suppresses oxidant damage to the
intestinal epithelium, and interferes with the formation of
BIOACTIVES AND TOXINS | Bioactives: Antioxidants 279
ADPFeoxygen complexes that initiate lipid peroxidation.
Phytic acid may play a role in the prevention of various cancers
including colon cancer.
Lignins and Lignans
Lignins range from 0.9% to 2.8% in whole-grain wheat, 1.9% to
5.6% in wheat bran, and 1.5% in wheat germ. They have been
shown to have antioxidant effects on ex vivo fresh lymphocytes.
They account for 2632% of the enterolactone (a mammalian
lignan) formed from cereal bran. Mammalian lignans are anti-
oxidants in vitro at concentrations of 10100 mM achievable
in vivo, particularly in the colon. Lignans make up 0.206,
2.86.7, and 0.49 mg/100 g of whole-grain wheat, wheat bran,
and wheat germ, respectively (Table 1). Lignins are neither
absorbed nor fermented in the human GIT. Only 04% of
lignins are apparently absorbed in rats following the consump-
tion of raw and processed wheat bran. Their antioxidant activity
due to the phenolic hydroxyl group may contribute to the
prevention of colon cancer.
Lipotropes and Methyl Donors (Betaine, Choline, and Folates)
Whole-grain wheat, wheat bran, and wheat germ contain about
0.28%, 1.04%, and 1.09% of betaine and choline, respectively,
and about 51, 231, and 420 mg folates/100 g. Betaine, choline,
and folates are lipotropic compounds together with methionine
and myoinositol that prevent excess fat deposition in the liver by
hastening fat removal or by limiting lipid synthesis. Whole-grain
wheat, wheat bran, and wheat germ can supply 0.51, 1.31, and
1.59 g lipotropes/100 g, respectively. Betaine is likely to be more
bioavailable as it is not bound to cell wall constituents as is the
case with ferulic acid. Data on apparent absorption of betaine
and choline are not available.
g-Oryzanols
g-Oryzanols or phytosteryl ferulates mainly found in rice bran
have been reported to have antioxidant activity. They are con-
sidered more effective than tocopherols and tocotrienols in the
prevention of cholesterol oxidation. They have the ability to
scavenge radicals in multiphase lipid systems and inhibit UV
irradiation-initiated linoleic acid oxidation.
Avenanthramides in Oats
Avenanthramides are found only in oats, particularly in the
outer layers. Avenanthramides or hydroxycinnamoyl anthrani-
linate alkaloids are amides of various cinnamic acids with dif-
ferent anthranilic acids. Avenanthramides 2p, 2f, and 2c are the
most abundant among over 35 forms identified to date. Oat
grain contains 413 mg avenanthramides 1/100 g (the major
avenanthramide), while oat bran contains 1.312.5 mg/100 g
according to the type of avenanthramide considered.
The inclusion of avenanthramide-2c (N-(30 ,40 -dihydroxy-
cinnamoyl)-5-hydroxyanthranilic acid) in rat diets altered
oxidantantioxidant balance in various tissues. It resulted in
selective reduction in the production of exercise-induced reac-
tive oxygen species and lipid peroxidation in the rats. This is
related to the ability of the avenanthramides to modulate
280 BIOACTIVES AND TOXINS | Bioactives: Antioxidants
antioxidant enzyme systems such as superoxide dismutase and
GSH reductase activities.
Saponins in Oats
Saponins are glycosides with a steroid or triterpenoid aglycone.
Oats synthesize two families of saponins, the steroidal avena-
cosides and the triterpenoid avenacins. The saponin content
varies from 0.02% to 0.13% (DW) in oat endosperm. Saponins
exert their antioxidant effects through the activation of tran-
scriptional activity of CuZn superoxide dismutase gene, scav-
enging of superoxide radicals, and reduction of lipid
peroxidation.
Phenolic Compounds
Perez-Jimenez et al. indicated 29 phenolic acids, 33 flavonoids,
6 lignans, and 2 other polyphenols as being present in various
cereal grains and cereal products.
Phenolic acids
Using the Phenol-Explorer database, Li and Beta identified
cereal and cereal products in which high maximum values
(>100 mg/100 g) have been reported for the sum of individual
phenolic acids. The list includes whole-grain maize, refined
maize flour, hard wheat semolina, common wheat germ,
whole-grain rye flour and bread, hard wheat whole-grain
flour, common wheat whole-grain flour, whole-grain oat
flour, refined rye flour, and whole-grain rice.
Phenolic acids occur as hydroxylated derivatives of benzoic
and cinnamic acids. The hydroxycinnamic acids are more com-
mon than hydroxybenzoic acids. Hydroxycinnamates identi-
fied in grains include ferulic, caffeic, p-coumaric, and sinapic
acids. Ferulic acid is the most abundant cinnamic acid deriva-
tive in cereal grains. Dehydrodimers of ferulic acid also exist
including 5-80 -dehydrodiferulic, 8-O-40 -dehydrodiferulic, and
5-80 -benzofuran dehydrodiferulic acids. Phenolic acids occur
as free, conjugated (soluble), and bound (insoluble) forms
with concentrations in the order of bound > conjugated > free.
The bound form makes up more than 80% of the total pheno-
lic acids. Phenolic acid composition is similar, but concentra-
tions of individual phenolic acids vary significantly in grain
botanical fractions of corn, wheat, oat, and barley. Vanillic,
p-coumaric, ferulic, sinapic, and syringic acids are present in all
grain fractions. Phenolic acids are unevenly distributed even
within the grain fractions of pericarp, aleurone, germ, and
endosperm obtained from the same cereal kernels.
Ferulic acid levels in whole-grain wheat, wheat bran, and
wheat germ are 16213, 138631, and 7124 mg/100 g,
respectively (Table 1). While absorption of ferulic acid is very
low following the consumption of whole-grain wheat or wheat
bran, 2.05.7% urinary excretion has been reported in humans
and rats. Small intestinal esterases may release very small
amounts of the bound ferulic acid. Absorption of free and
conjugated ferulic acid is reported as high in the rat small
intestine; however, 27.878.9% is excreted through the urine
in humans. Insoluble fiber plays a role in delivering
antioxidant-bound phenolics to the colon thereby potentially
aiding in the prevention of colon cancer and contributing to
gut health.
Flavonoids
Using the Phenol-Explorer database, Li and Beta reported the
sum of individual flavonoids (mg/100 g) of selected cereals
and cereal products as 51.5108.4 for common wheat whole-
grain flour, 16.463.4 for whole-grain buckwheat flour,
32.739.0 for whole-grain barley flour, 13.226.3 for common
refined wheat flour, 1.717.0 for refined buckwheat flour, and
4.013.4 for buckwheat thermally treated groats. The major
flavonoids include apigenin arabinosideglucoside and api-
genin galactosidearabinoside in common wheat whole-grain
flour; quercetin 3-O-rutinoside and apigenin 6-C-glucoside in
whole-grain buckwheat flour; prodelphinidin dimer B3, pro-
cyanidin dimer B3, and ()-catechin in whole-grain barley
flour; apigenin galactosidearabinoside and apigenin
arabinosideglucoside in common refined wheat flour; and
quercetin 3-O-rutinoside in refined buckwheat flour.
Proanthocyanidin polyphenols with degree of polymerization
(dp) greater than 10 (1307.31767.0 mg/100 g), dp 710
(150.0325.0), and dp 46 (85.1253) are present in some
varieties of whole-grain sorghum.
Due to their antioxidant properties, phenolic compounds
are believed to potentially play a role in protecting the body
against oxidative stress and its effects. Free and bound phenolic
compounds in grains may directly mitigate oxidative stress by
increasing plasma total antioxidant capacity or do so indirectly
through cell signaling. It is suggested that phenolic compounds
can modulate the GSH radical scavenging system to improve
plasma total antioxidant capacity. In this radical scavenging
system, GSH peroxidase is used to metabolize hydrogen per-
oxide to water by using reduced GSH as a hydrogen donor. The
ability of reduced GSH to scavenge free radicals can be com-
promised if oxidized GSH is not converted back to GSH by
GSH reductase or if the activity of GSH peroxidase is reduced.
Some in vitro evidence suggests that flavonoids can alter the
expression of genes responsible for the synthesis and regula-
tion of GSH. Furthermore, it is reported that flavonoids can
increase plasma total antioxidant capacity by increasing
plasma uric acid levels, which has reducing and free radical
scavenging activities.
Alkyresorcinols
Alkyresorcinols are found in higher concentration in rye
(320 mg/100 g) compared to wheat (12129 mg/100 g)
(Table 1). The phenolic lipids consist of a phenolic ring with
two hydroxyl groups and long (typically C15C25) non-
isoprenoid side chain attached at the C5 position. The major
alkylresorcinols include 5-heneicosylresorcinol, 5-heneicosenyl-
resorcinol, 5-nonadecylresorcinol, and 5-heptadecylresorcinol
in rye and wheat products. About 6079% of alkylresorcinols
are absorbed in the small intestine of pigs following the con-
sumption of whole-grain rye bread. The 1,3-dihydroxybenzene
derivatives are absorbed (4571%) within the small intestine of
ileostomates following the consumption of baked rye products.
They are considered potential biomarkers of whole-grain rye
and wheat intake for epidemiological research and observa-
tional studies. Although they are known to exert antioxidant
effects through the modulation of lipid oxidation, their biolog-
ical activity is multifactorial.

Carotenoids
Carotenoids found in grains include beta-carotene, beta-
cryptoxanthin, lutein, and zeaxanthin. Table 1 shows the
total carotenoid, beta-carotene, lutein, zeaxanthin, and beta-
cryptoxanthin content of whole-grain wheat and wheat bran.
Only beta-carotene content of wheat germ is included.
Although the composition of carotenoids is similar among
cereals (wheat, oat, corn, and barley), concentrations vary
significantly. Total carotenoid content ranges are 0.882.27
and 14.1731.35 mg kg
1, respectively, in corn and non-corn
(wheat, oat, and barley) cereal endosperm fractions. The aleu-
rone layer, which botanically is the outer part of the endo-
sperm, has zeaxanthin contents two- to fivefold higher than
lutein. Lutein and zeaxanthin contents are higher in the germ
of non-corn cereals than in corn germ. The germ is the most
concentrated source of carotenoids in cereals. There is a
high correlation between %DPPH free radical scavenging and
total lutein and zeaxanthin content. Beta-carotene, beta-
cryptoxanthin, lutein, and zeaxanthin exert antioxidant effects.
Lutein and zeaxanthin quench singlet oxygen, neutralize
photosensitizers, and inhibit lipid peroxidation.
Tocopherols and Tocotrienols
Tocopherols and tocotrienols (vitamin E vitamers) are lipid-
soluble micronutrients found in grains. Table 1 shows the
content of total tocols, tocopherols, and tocotrienols in
wheat. Through the exertion of their strong intracellular anti-
oxidant effects, tocols protect against oxidative damage of
polyunsaturated fatty acids within cell membranes and in lipo-
proteins, DNA nucleotidic bases, and proteins. The tocotrie-
nols are considered stronger antioxidants than tocopherols.
Policosanols
Policosanols are present at low levels in the germ of wheat
(1.01 mg/100 g), while wheat bran (0.113.00 mg/100 g) and
whole-grain wheat (0.305.62 mg/100 g) contain lower levels
(Table 1). They exert antioxidant effects through the reduction
of low-density lipoprotein and membrane lipid peroxidation.
Melatonin
Melatonin constitutes 0.20.4 mg/100 g of whole-grain wheat.
There is no data on melatonin absorption in humans following
the consumption of whole-grain wheat. Melatonin exerts its
antioxidant effects through several mechanisms. It increases
gene expression for antioxidant enzymes. It is reported to
protect against lenticular protein oxidation enzymes. Melato-
nin is considered the most potent physiological scavenger of
l
hydroxyl radical (OH ).
Future Prospects
The antioxidant effects of bioactive compounds from whole
grains are potentially beneficial in delaying or preventing the
development of diseases in relation to increased oxidative
stress. However, the same compounds that exert antioxidant
BIOACTIVES AND TOXINS | Bioactives: Antioxidants 281
effects have other physiological functions. Unraveling the
cumulative mechanisms involved beyond antioxidant effects
will provide strong evidence for consumption of whole grains
concentrated in bioactives.
Exercises for Revision
What are the major grain bioactives?
Explain briefly how bioactives can be altered through grain
processing.
What is meant by the term oxidative stress and what is its
significance in relation to chronic diseases of lifestyle?
Which phenolic acid is predominant in grains?
How are phenolic compounds purported to protect the
body against oxidative stress and its effects?
Briefly explain how minerals in grains serve as antioxidants.
What are the major carotenoids in grains?
Exercises for Readers to Explore the Topic Further
The content of major phytochemicals such as phenolic acids
and flavonoids may be high in grains; however, these anti-
oxidants are tightly bound to the cell wall matrix. Explore the
extent of their bioavailability taking into consideration those
studies that have shown harmful effects on human health of
some antioxidants under certain conditions.
Several approaches are used to assess antioxidant capacity
of bioactive components in grains. Discuss the limitations
of chemical- and cellular-based models in establishing the
optimum doses to be taken daily to achieve beneficial
effects on human health.
There is a need for ex vivo and in vivo studies to confirm the
efficacy and site of action of grain bioactives in animals
including humans. Investigate a selected bioactive in
depth, and propose what further detailed studies would
be required to achieve a fuller understanding of its action.
Synergistic mechanisms and modes of action among bio-
actives and with other grain components in whole-grain
products are little understood. Moreover, bioactives may
exert their effects through mechanisms other than antioxi-
dant action. Can you suggest general approaches to identify
suitable biomarkers to conduct such mechanistic studies in
a holistic manner?
See also: Bioactives and Toxins: Bioactive Compounds in Wheat
Bran; Food Grains and the Consumer: Grains and Health; Food
Grains and Well-being: Functional Foods: Dietary Fibers,
Prebiotics, Probiotics, and Synbiotics; Functional Foods: Overview;
The Basics: The Grain Crops: An Overview; The Cereal Grains: An
Overview of the Family of Cereal Grains Prominent in World
Agriculture.
Further Reading
Adom KK, Sorrells ME, and Liu RH (2005) Phytochemicals and antioxidant activity of
milled fractions of different wheat varieties. Journal of Agricultural and Food
Chemistry 53: 663671.
282 BIOACTIVES AND TOXINS | Bioactives: Antioxidants
Aruoma OI (1998) Free radicals, oxidative stress, and antioxidants in human health and
disease. Journal of the American Oil Chemists Society 75: 199212.
Belodrajdic DP and Bird AR (2013) The potential role of phytochemicals in wholegrain
cereals for the prevention of type-2 diabetes. Nutrition Journal 12: 62.
Fardet A (2010) New hypothesis for the health-protective mechanisms of
whole-grain cereals: What is beyond fibre? Nutrition Research Reviews
23: 65134.
Grassi D, Desideri G, and Ferri C (2010) Flavonoids: Antioxidants against
atherosclerosis. Nutrients 2: 889902.
Li W and Beta T (2013) Food sources of phenolics compounds. In: Ramawat KG and
Merillon JM (eds.) Natural Products: Phytochemistry, Botany and Metabolism
of Alkaloids, Phenolics and Terpenes, pp. 25272558. Berlin, Heidelberg:
Springer-Verlag.
Ndolo VU and Beta T (2013) Distribution of carotenoids in endosperm, germ, and
aleurone fractions of cereal grain kernels. Food Chemistry 139: 663671.
Ndolo VU and Beta T (2014) Comparative studies on composition and
distribution of phenolic acids in cereal grain botanical fractions. Cereal Chemistry
91: 522530.
Neveu V, Perez-Jimenez J, Vos F, et al. (2010) Phenol-Explorer: An online
comprehensive database on polyphenol contents in foods. Database. doi:10.1093/
database/bap024. Version 1.5.2. http://www.phenol-explorer.eu.
Okarter N and Liu RH (2010) Health benefits of whole grain phytochemicals. Critical
Reviews in Food Science and Nutrition 50: 193208.
Perez-Jimenez J, Neveu V, Vos F, and Scalbert A (2010) Systematic analysis of the
content of 502 polyphenols in 452 foods and beverages: An application of the
Phenol-Explorer database. Journal of Agricultural and Food Chemistry
58: 49594969.
Ryan D, Kendall M, and Robards K (2007) Bioactivity of oats as it relates to
cardiovascular disease. Nutrition Research Reviews 20: 147162.
 The Antinutritional Components of Grains
CK Madsen and H Brinch-Pedersen, Aarhus University, Slagelse, Denmark
2016 Elsevier Ltd. All rights reserved.
Topic Highlights
Antinutritional compounds interfere with the utilization of
nutrients and energy.
The major antinutritionals of grains are phytate, proteina-
ceous inhibitors of digestive enzymes, tannins, and non-
starch polysaccharides.
Antinutritionals may also offer nutritional benefits that
have to be balanced against their negative effects.
The significance of antinutritionals depends on species and
nutritional status of the human or animal consuming it.
Learning Objectives
To achieve an understanding of the chemistry, occurrence,
and antinutritional mechanism of the major antinutritional
compounds in grains.
To gain an overview of strategies to counter
antinutritionals.
Introduction
Antinutritionals are compounds that reduce the bioavailability
of essential nutrients or energy in the diet. This leads to dis-
crepancies between the formal and actual nutritional values of
a food or feed. It is therefore most important that both nutri-
ents and antinutritionals are part of the equation when the
nutritional value of a food item is evaluated. Moreover, it is
important to notice that some antinutritionals may also pos-
sess positive dietary properties and that nutritional goals in
human and animal nutrition can be very different, that is,
with respect to weight gain. Antinutritionals that are deleteri-
ous for the calorie yield of a diet may be used to counter
obesity in a human diet, whereas they result in poor resource
use efficiency in animal husbandry.
The use of the term antinutritionals is inconsistent in the
scientific literature and may sometimes refer to compounds
that are considered overall unhealthy (i.e., allergens, irritants,
and toxins). Such compounds are beyond the scope of this
article.
Phytate
Phytic acid or myo-inositol-(1,2,3,4,5,6)hexakisphosphate
(IP6) is the major phosphorus (P) storage compound in
plants. IP6 is hexaphosphorylated myo-inositol. At physiolog-
ical pH, it adopts a conformation with two phosphates in the
negatively charged phosphate groups (Figure 1). The strength
of the interactions depends on the charge of the cations
(Me3 > Me2 > Me) and to a lesser extent other parameters
such as ionic radius. At acidic pH, the metal cations compete
with H and interactions with most metal cations are insignif-
icant below pH 5.
In grains, IP6 typically accounts for 2/3 of the total P
content (see Table 1 for exact values). This IP6 is found as an
insoluble mixed salt with cations called phytin. In many grains
including cereals and legumes, phytin forms spherical inclu-
sions called globoids inside protein storage vacuoles. Not only
are the globoids the principal site of P, K, and Mg in the mature
cereal grain, but also they contain Ca, Fe, Zn, Cu, Mn, and Na.
Globoids are predominantly found in the aleurone layer of
small grain cereals and in the embryo of maize, tissues that
are the main contributors to millers bran fraction. Accord-
ingly, the IP6 contents in bran and white flour are higher and
lower, respectively, than those in whole grain. IP6 is resistant to
most phosphatases, whereas the lower inositol phosphates are
substrates of a wider range of phosphatases. Phosphatases that
can initiate dephosphorylation of IP6 are known as phytases.
All plants express phytases to assist in their IP6 metabolism but
the phytase activity in mature seed varies several orders of
magnitude, even in closely related species. This is in sharp
contrast to the modest variation in total and proportional
content of IP6 (Table 1). The Triticeae tribe of cereals (such
as wheat, barley, rye, and triticale) stand out with exceptionally
high mature grain phytase activity. This is caused by the rela-
tively recent duplication of a purple acid phosphatase phytase
gene whose archaic function was to be expressed during ger-
mination. Following the gene duplication, one copy has chan-
ged expression pattern so it is expressed during grain filling.
The reported intake of IP6 for adults ranges from 180 to
5577 mg day1. Omnivorous Westerners are in the low range
of the interval (usually less than 1000 mg day1). Vegetarians
and people in developing countries are in the high range
(>1000 mg day1). The highest value (5577 mg day1) has
been reported for a group of Trappist monks on a lacto-ovo
vegetarian diet. The utilization of IP6 by humans and most
nonruminants is negligible unless the meal has sufficient phy-
tase activity. Fast-growing animals on a high phytate contain-
ing feed without P supplements may suffer from P deficiency.
The low retention of IP6 affects not only the molecule per se
but also its chelation partners. Because of the chemical prop-
erties discussed earlier in the text, IP6 will release chelated
metals in the low pH of the stomach but recruit ions again
from the food matrix as pH reaches neutral during the duode-
nal passage. Therefore, IP6 from one source exerts an antinu-
tritional effect on the entire meal. In vivo, this has been shown
to affect zinc, iron, calcium, magnesium, copper, and manga-
ne