Sie sind auf Seite 1von 664


Postharvest Ripening
Physiology of Crops

Edited by
Sunil Pareek
Postharvest Ripening
Physiology of Crops
Series Editor
Sunil Pareek
Department of Agriculture and Environmental Sciences
National Institute of Food Technology Entrepreneurship and Management
Kundli, Sonepat, Haryana, India

Postharvest Ripening Physiology of Crops (2016)

Edited by Sunil Pareek
Postharvest Ripening
Physiology of Crops
Edited by
Sunil Pareek

Boca Raton London New York

CRC Press is an imprint of the

Taylor & Francis Group, an informa business
CRC Press
Taylor & Francis Group
6000 Broken Sound Parkway NW, Suite 300
Boca Raton, FL 33487-2742
© 2016 by Taylor & Francis Group, LLC
CRC Press is an imprint of Taylor & Francis Group, an Informa business

No claim to original U.S. Government works

Version Date: 20151207

International Standard Book Number-13: 978-1-4987-0381-9 (eBook - PDF)

This book contains information obtained from authentic and highly regarded sources. Reasonable
efforts have been made to publish reliable data and information, but the author and publisher cannot
assume responsibility for the validity of all materials or the consequences of their use. The authors and
publishers have attempted to trace the copyright holders of all material reproduced in this publication
and apologize to copyright holders if permission to publish in this form has not been obtained. If any
copyright material has not been acknowledged please write and let us know so we may rectify in any
future reprint.

Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced,
transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or
hereafter invented, including photocopying, microfilming, and recording, or in any information stor-
age or retrieval system, without written permission from the publishers.

For permission to photocopy or use material electronically from this work, please access www.copy- ( or contact the Copyright Clearance Center, Inc. (CCC), 222
Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organization that pro-
vides licenses and registration for a variety of users. For organizations that have been granted a photo-
copy license by the CCC, a separate system of payment has been arranged.

Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are
used only for identification and explanation without intent to infringe.
Visit the Taylor & Francis Web site at
and the CRC Press Web site at


1 Ripening Physiology: An Overview 1

Abstract 2
1.1 Introduction 2
1.2 Climacteric Phenomenon 4
1.3 Physicochemical and Metabolic Changes 12
1.3.1 Color Changes 13
1.3.2 Sugar Changes 14
1.3.3 Organic Acid Changes 16
1.3.4 Flavor and Aroma Changes 18
1.3.5 Cell Wall and Textural Changes 22
1.3.6 Physiological Changes 28
1.4 Conclusions and Future Perspectives 33
References 33

2 Postharvest Physiology of Fruits and Vegetables 49

Abstract 50
2.1 Introduction 51

Co ntents

2.2 Classification of Fruits and Vegetables Based on

Physiological Characteristics 51
2.2.1 Ethylene Biology 51
2.2.2 Respiratory Characteristics: Climacteric and
Nonclimacteric 56
2.2.3 Developmental or Maturity Stage at Harvest 56
2.2.4 Tolerance to Low Temperatures 58
2.3 Factors Affecting Physiology of Fruits and Vegetables in
Postharvest Systems 60
2.3.1 Temperature 60
2.3.2 Humidity 62
2.3.3 Atmospheric Modification 64
2.3.4 Abiotic Stresses 65
2.4 Physiological Changes Occurring during Postharvest
Handling or Storage 66
2.4.1 Depletion of Respiratory Substrate 66
2.4.2 Hormonal Effects 68
2.4.3 Membrane Alterations 71
2.5 Conclusions and Future Perspectives 71
References 72

3 Postharvest Quality of Ornamental Plants 81

Abstract 82
3.1 Introduction 82
3.2 Quality Attributes in Ornamental Plants 83
3.3 Influence of Water Relations on Ornamental Longevity 85
3.4 Action of Ethylene on Ornamental Plant Quality 88
3.4.1 Inhibition of Ethylene Synthesis 91
3.4.2 Inhibition of Ethylene Action 93
3.4.3 Ethylene Absorbers 95
3.5 Role of Abscisic Acid, Gibberellins, and Cytokinins 96
3.6 Role of Calcium on Flower Senescence 97
3.7 Respiration 98

Co ntents

3.8 Temperature 99
3.9 Handling of Cut Flowers 102
3.10 Potted Plants 103
3.11 Conclusions and Future Perspectives 104
Acknowledgment 105
References 105

4 Physiology and Molecular Biology of

Flower Senescence 109
Abstract 110
4.1 Introduction 110
4.2 Ethylene and Senescence of Cut Flowers 111
4.2.1 Ethylene Response of Cut Flowers 111
4.2.2 Types of Senescence in Cut Flowers with High
Ethylene Sensitivity 112
4.2.3 Ethylene in Petal-Wilting-Type Flowers 113
4.2.4 Ethylene in Petal-Abscission-Type Flowers 114
4.2.5 Ethylene Biosynthesis 114
4.2.6 Ethylene Signal Transduction 115
4.2.7 Acceleration of Flower Senescence by
Pollination 116
4.2.8 Acceleration of Flower Senescence by
Wounding 119
4.2.9 Effect of Temperature on Ethylene Production
and Perception 119
4.3 Plant Hormones Other than Ethylene in Flower
Senescence 120
4.3.1 Auxin 120
4.3.2 Gibberellin 120
4.3.3 Cytokinin 120
4.3.4 Abscisic Acid 121
4.3.5 Jasmonic Acid 122
4.4 Programmed Cell Death in Flower Senescence 122
4.4.1 Programmed Cell Death 122

Co ntents

4.4.2 Gene Expression during PCD in Flowers 123

4.4.3 Autophagy in Petal Senescence 124
4.5 Conclusions and Future Perspectives 124
References 125

5 Respiratory Metabolism 139

Abstract 140
5.1 Introduction 140
5.2 Why Measure Respiration? 141
5.3 Major Components of Respiration 142
5.3.1 Glycolysis 142
5.3.2 Pentose-Phosphate Shunt 142
5.3.3 Anaerobic Diversion 143
5.3.4 Tricarboxylic Acid Cycle 143
5.3.5 Electron Transport (Chemiosmotic
Phosphorylation) 145
5.4 Measurement of Respiration 146
5.4.1 Loss of Substrate, Heat Production, and Water 146
5.4.2 Consumption of Oxygen and Production of
Carbon Dioxide 147 Static System 147 Flow-Through or Dynamic System 149
5.5 Sampling and Analyzing 151
5.6 Instruments and Techniques 152
5.7 Pre- and Postharvest Factors Affecting Respiration 152
5.7.1 Temperature Effects 152
5.7.2 Respiratory Quotient 154
5.7.3 Physical Stress 154
5.7.4 Internal Factors 155 Genotype 155 Type of Plant Part 155 Respiratory Climacteric 155
5.8 Conclusions and Future Perspectives 156
References 156

Co ntents

6 Stomata and Postharvest Physiology 157

Abstract 158
6.1 Introduction 158
6.2 Stomata 159
6.2.1 Role of Stomata in Plants 159
6.2.2 Mechanism of Stomatal Closure and Opening 161
6.2.3 Signal Transduction Pathways in Guard Cells
for Stomatal Closure 164
6.3 Role of Stomata in the Postharvest Phase 166
6.3.1 Stomata in Relation to Vase Life of Cut Flowers 166
6.3.2 Stomata in Relation to Quality of Vegetables 169
6.3.3 Stomata in Relation to Quality of Fruits 172
6.4 Preharvest Conditions Leading to Postharvest Problems
via Stomata 174
6.4.1 Relative Humidity and Stomata Control 175 How Does Preharvest Low VPD
Affect Postharvest Stomata Control? 176 Induction of Stomata Morphological
Changes by Low VPD 179
6.4.2 Temperature 181
6.4.3 Light 182
6.5 Stomata and Tolerance to Postharvest Diseases and
Physiological Disorders 184
6.6 Postharvest Treatments and Stomata 186
6.7 Ethylene, Stomata, and Senescence 187
6.8 Conclusions and Future Perspectives 190
References 191

7 Water Loss from Harvested Horticultural

Commodities 217
Abstract 218
7.1 Importance of Water Loss 219
7.2 Properties of Water 220

Co ntents

7.3 Psychrometrics: Behavior of Water in Air 221

7.4 Transpiration: Diffusion of Water Vapor 224
7.5 Resistance to Diffusion of Water Vapor 226
7.6 Measurement of Transpiration 226
7.6.1 Weight Loss 227
7.6.2 Direct Measurement 227
7.6.3 Diffusion Porometer 227
7.7 Factors Affecting Water Loss 227
7.7.1 Commodity Factors 227 Surface-to-Volume Ratio 228 Routes of Water Loss 228 Anatomy of the Evaporating Surface 229 Physiological State of the
Commodity 230 Cultivar 230 Cultural Conditions 231
7.7.2 Environmental Factors 231 Humidity 231 Diffusion Shells and Air Velocity 231 Temperature 232 Atmospheric Pressure 232
7.8 Methods to Reduce Water Loss 232
7.8.1 Handling Techniques 232
7.8.2 Proper Refrigeration Design 232
7.8.3 Packaging 233
7.8.4 Waxing 234
7.8.5 Film Wraps 234
7.8.6 Curing 234
7.9 Conclusions and Future Perspectives 235
References 235

8 Lysophospholipids and Postharvest Quality

of Fruits, Vegetables, and Cut Flowers 237
Abstract 238
8.1 Introduction 238

Co ntents

8.2 Lipids as Signal Molecules in Plant Senescence and

Stress Response 239
8.2.1 Chemistry 239
8.2.2 Metabolism 240
8.3 Modulation of Postharvest Quality by Lysophospholipids 241
8.3.1 Effects on Color 242
8.3.2 Effects on Texture 242
8.3.3 Other Effects on Postharvest Quality-Related
Features 246
8.4 LPE Treatment Condition 247
8.5 Improvement of Postharvest Quality by
Lysophospholipids: Potential and Limitations 248
8.6 Conclusions and Future Perspectives 249
References 249

9 Fruit Skin Color and the Role of Pigments

during Fruit Ripening 255
Abstract 256
9.1 Introduction 256
9.2 Economics of Color Fruit 258
9.3 Carotenoid Pigments in Fruits 259
9.4 Anthocyanin Pigments in Fruits 260
9.5 Grape as a Model System for Pigmentation Studies
in Fruit Crops 270
9.6 Case Studies 278
9.6.1 Apple 278
9.6.2 Strawberry 279
9.6.3 Tomato 280
9.6.4 Litchi 281
9.6.5 Chinese Bayberry 282
9.6.6 ‘Hass’ Avocados 283
9.6.7 Pear 283
9.6.8 Cherry 284
9.6.9 Kiwifruit 284

Co ntents

9.7 Pigments from Fruit to Human Health 284

9.8 Transgenic Plants Developed for Fruit Color 285
9.9 Conclusions and Future Perspectives 286
References 288

10 Molecular Regulation of Fruit Ripening 299

Abstract 300
10.1 Introduction 300
10.2 Transcriptional Control of Fruit Ripening 301
10.3 Hormonal and Transcriptional Regulation during
Ripening 305
10.3.1 Ethylene 306
10.3.2 Auxins 308
10.3.3 Gibberellins 308
10.3.4 Abscisic Acid 308
10.4 Epigenetic Regulation of Fruit Development and
Ripening 309
10.5 Ripening Pathways and Associated Fruit Quality 312
10.5.1 Sugar Accumulation 313
10.5.2 Cell Walls and Fruit Shelf Life 313
10.5.3 Color, Flavor, and Nutrition in Fruits 314 Chloroplast-to-Chromoplast
Conversion 314 Flavor and Aroma Production 315 Control of Color and Texture
Changes 316
10.6 Human Nutrition and Fruits 317
10.6.1 Human Nutrition: Functional Genomics/
Systems Approach 320
10.7 Horticultural Crop Improvement 321
10.8 Genetic Manipulation of Ripening Regulatory
Genes 322
10.9 Conclusions and Future Perspectives 322
References 324

Co ntents

11 Advances in Ethylene Signal Transduction in

Fruits and Vegetables 339
Abstract 340
11.1 Introduction 340
11.2 Ethylene Physiology in Climacteric and Nonclimacteric
Fruits 341
11.3 Ethylene Signaling Components in Fruits 342
11.4 Analyses of Ethylene Signal Components in Other Fruit
Species 344
11.4.1 Climacteric Fruits 344
11.4.2 Nonclimacteric Fruits 348
11.5 Transcriptional Regulators of Fruit Ripening 349
11.6 Conclusions and Future Perspectives 352
References 353

12 Internal Atmosphere of Fruits: Role and

Significance in Ripening and Storability 359
Abstract 360
12.1 Introduction 361
12.2 Endogenous Volatiles in Fruits 363
12.3 Factors Affecting and Basis of Internal Atmosphere
of Harvested Fruit 363
12.4 Variability in the Internal Atmosphere of Fruits 366
12.5 Influence of the Internal Atmosphere on Ripening
and Related Aspects 367
12.5.1 Ripening 367
12.5.2 Flavor and Aroma 368
12.5.3 Fruit Decay 370
12.6 Ripening Behavior of Some Fruits under Attached
and Detached Conditions 371
12.6.1 Tomato 371
12.6.2 Melons 372
12.6.3 Sweet Pepper 372
12.6.4 Saskatoon 373

Co ntents

12.7 Role of Some Gases and Endogenous Volatiles in

Fruit Ripening 373
12.7.1 Ethylene 373 Role of Ethylene in Ripening of
Climacteric Fruits 373 Role of Ethylene in Ripening of
Some Nonclimacteric Fruits 375
12.7.2 Oxygen and Carbon Dioxide 379 Low Oxygen 379 High Carbon Dioxide 382 Ratio of O2 to CO2 383
12.7.3 Ethanol and Acetaldehyde 384
12.7.4 Water Vapors and Water Status in Fruit 385
12.7.5 Salicylic Acid and Methyl Salicylate 387
12.7.6 Jasmonic Acid and Jasmonates 387
12.7.7 Nitric Oxide 388
12.8 Internal Atmosphere of Fruits: Practical Implication
in Ripening and Storability 388
12.9 Conclusions and Future Perspectives 390
References 391

13 Proteomics of Fruit Development and

Ripening 413
Abstract 414
13.1 Introduction 414
13.2 Characteristics of Climacteric and Nonclimacteric
Fruit Ripening 416
13.3 Proteomic Tools and Approaches for Fruit Ripening
Assessment 417
13.3.1 Challenges in Proteomics Analysis 418
13.3.2 Protein Sample Preparation: Tissue Disruption,
Homogenization, and Solubilization 418
13.3.3 Protein Separation: Gel-Based and Gel-Free
Technologies 421

Co ntents

13.4 Proteomes of Climacteric and Nonclimacteric Fruits 422

13.4.1 Overrepresented Functional Categories
during Development and Ripening 422
13.5 Pathway Analysis Using KEGG 426
13.5.1 Proteins Associated with Carbohydrate
Metabolism 426
13.5.2 Proteins Associated with Energy Metabolism 431 Proteins with a Role in Carbon
Fixation 431 Proteins Associated with the
Pentose Phosphate, Glycolysis,
and Pyruvate Metabolisms 432 Proteins Involved in the
TCA Cycle 436
13.5.3 Proteins Involved in Amino Acid Metabolism
and Ethylene Biosynthesis 436
13.5.4 Proteins Involved in Flavonoid Biosynthesis 437
13.6 Conclusions and Future Perspectives 438
References 439

14 Potato Tuber Dormancy and Postharvest

Sprout Control 449
Abstract 450
14.1 Introduction 451
14.2 Dormancy: General Considerations 451
14.2.1 Developmental Aspects 452
14.3 Genetics of Tuber Dormancy 453
14.4 Pre- and Postharvest Environmental Effects 453
14.5 Physiological Regulation of Tuber Dormancy 454
14.5.1 Auxin 455
14.5.2 Carotenoid-Derived Hormones:
Abscisic Acid and Strigolactone 456
14.5.3 Cytokinins 458
14.5.4 Ethylene 459

Co ntents

14.5.5 Gibberellins 460

14.5.6 Summary and Conclusions 460
14.6 Transcriptional Regulation during Dormancy 461
14.6.1 Initiation of Tuber Dormancy 461
14.6.2 Dormancy Termination 462
14.6.3 Epigenetic Control of Tuber Dormancy 463
14.7 Control of Sprouting in Storage 463
14.7.1 Introduction and Importance 463
14.7.2 Dormancy, Cultivar Selection, and
Storage Temperature 464
14.7.3 Chemical Control 464
14.8 Conclusions and Future Perspectives 467
References 467

15 Calcium Deficiency Disorders in Plants 477

Abstract 478
15.1 History of Ca2+ Deficiency Disorders 479
15.2 Role of Ca as an Essential Plant Macronutrient
2+ 479
15.3 Symptoms of Ca2+ Deficiency Disorders
in Fruit 481
15.3.1 Apple 481
15.3.2 Tomato 484
15.3.3 Watermelon 484
15.3.4 Pepper 484
15.4 Symptoms of Ca2+ Deficiency Disorders in
Leafy Vegetables 485
15.4.1 Lettuce 485
15.4.2 Cauliflower 485
15.4.3 Artichoke 486
15.4.4 Celery 487
15.5 Potential Mechanisms Regulating Ca2+ Deficiency
Disorders 488
15.5.1 Total Tissue Ca2+ Content 488

Co ntents

15.5.2 Cellular Regulation of Ca2+ Partitioning

and Distribution 490
15.5.3 Other Nutrients 492 Nitrogen 492 Potassium and Magnesium 493 Boron 494 Phosphorus 494
15.5.4 Reactive Oxygen Species 495
15.5.5 Growth Regulators 495 Growth Regulators Affecting
Total Tissue Ca2+ Content 496 Growth Regulators Influencing
Cellular Ca2+ Distribution 498 Growth Regulator Effect on
Oxidative Metabolism 499
15.6 Possible Control Strategies 500
15.6.1 At the Tissue Level 500
15.6.2 At the Cellular Level 501
15.7 Final Considerations and Future Research
Needs 501
References 502

16 Fresh Fruit Aroma: An Integrative Overview

for a Complex Flavor Trait 513
Abstract 514
16.1 Introduction 515
16.2 Aroma Composition in Fruits 516
16.2.1 Apple 516
16.2.2 Melon 517
16.2.3 Strawberry 517
16.2.4 Tomato 518
16.2.5 Citrus 518
16.2.6 Grape 519

Co ntents

16.2.7 Peach 519

16.2.8 Banana 520
16.3 Biosynthesis and Regulation of Aroma Volatiles in Fruit 520
16.3.1 Biosynthetic Pathways of Aroma Volatiles 520 Fatty Acid Metabolism 520 Amino Acid Metabolism 522 Ester Biosynthesis 522 Carbohydrate Metabolism 524
16.3.2 Aroma Modulation during Fruit Ripening 525
16.4 Influence of Pre- and Postharvest Factors on Fruit
Aroma 527
16.4.1 Preharvest Factors 527 Genotype 527 Growing Conditions 529 Fruit Maturity 531
16.4.2 Postharvest Technologies 532 Storage Temperature 532 Storage Atmosphere 534 Ethylene Control 536 Other Technologies 537
16.5 Conclusions and Future Perspectives 538
Acknowledgments 539
References 539

17 Flavor and Aroma Compounds of Some

Exotic Tropical Fruits and Berries: Biosynthetic
Pathways and Metabolism 553
Abstract 554
17.1 Introduction 555
17.2 Exotic Fruits and Their Flavor Profiles 556
17.2.1 Lychee (Litchi chinensis) 556
17.2.2 Rambutan (Nephelium lappaceum L.) 556
17.2.3 Yellow Passion (Passiflora edulis) 558
17.2.4 Durian Fruit (Durio zibethinus) 559

Co ntents

17.2.5 Star Fruit or Carambola (Averrhoa carambola L.) 560

17.2.6 Mangosteen (Garcinia mangostana) 560
17.2.7 Snake Fruit (Salacca edulis Reinw) 560
17.2.8 Costa Rican Guava (Psidium
friedrichsthalium) 565
17.2.9 Pitanga Fruit (Eugenia uniflora L.) 566
17.2.10 Umbu-Caja Fruit (Spondias citherea) 566
17.2.11 Camu-Camu Fruit (Myrciaria dubia) 566
17.2.12 Cupuacu Fruit (Theobroma grandiflorum) 567
17.2.13 Araca-Boi Fruit (Eugenia stipitata) 567
17.2.14 Mangaba Fruit (Hancornia speciosa
17.2.15 Garcinia Fruit (Garcinia dulcis Kurz) 568
17.2.16 Guabiju Fruit (Myrcianthes pungens Berg) 568
17.2.17 Guabiroba Fruit (Campomanesia xanthocarpa
17.2.18 Bacuri Fruit (Platonia sculenta) 568
17.2.19 Cashew Fruit (Anacardium occidentale L.) 569
17.2.20 Melon Fruit (Cucumis melo) 569
17.2.21 Jackfruit (Artocarpus heterophyllus Lam.) 570
17.2.22 Sapodilla Fruit (Achras sapota L.) 570
17.2.23 Genipap Fruit (Genipa americana) 571
17.2.24 Soursop Fruit (Annona muricata) 571
17.2.25 Acerola Fruit (Malphigia glabra L.) 572
17.2.26 Tamarind Fruit (Tamarindus indica L.) 572
17.2.27 Velvet Tamarind (Dialium guineense) 572
17.2.28  African Star Apple Fruit (Chrysophillum albidum) 573
17.3 Aroma Compounds’ Biosynthetic Pathways and
Metabolism 573
17.3.1 Fruit’s Volatile Ester Metabolism 573
17.3.2 Fruit’s Volatile Terpenoid Metabolism 575
17.3.3 Sulfur Volatile Compounds’ Biosynthetic
Pathway 577
17.4 Conclusions and Future Perspectives 579
References 579

Co ntents

18 Impact of Postharvest Technologies on the

Flavor of Fresh Fruits and Vegetables 585
Abstract 586
18.1 Introduction 587
18.2 Flavor of Fruits and Vegetables 588
18.2.1 Sensory Assessment 588
18.2.2 Chemical Flavor Constituents 589
18.3 Mechanism of Flavor Change 590
18.3.1 Metabolic Changes 591
18.3.2 Diffusional Changes 592
18.4 Impact of Postharvest Technologies 593
18.4.1 Ripening Manipulation 593
18.4.2 Temperature and Cold Storage 595
18.4.3 Controlled Atmosphere Storage 597
18.4.4 Packaging 599
18.4.5 Edible Coatings 601
18.4.6 Postharvest Treatments 601 Cutting 602 Heat Treatments 603 Irradiation 604 Ozone 605 Chemical Fumigation 606
18.5 Flavor Enhancement 607
18.6 Conclusions and Future Perspectives 607
References 608


To the Late Professor Adel A. Kader

During his more than 35 years at the University of California, Davis (UC
Davis), Professor Adel Kader maintained an extremely active teaching,
research, and extension program in postharvest biology and technology of
horticultural crops. His prodigious output includes more than 230 technical
manuscripts, many book chapters, and numerous extension publications. He
was the most cited author by postharvest colleagues. Kader was the undis-
puted leader of a research team that has reached high levels of knowledge
in the different disciplines; he was a reference for researchers, enterprises,
and international institutes. His research efforts were initially focused on
improving the more obvious aspects of the postharvest quality of fruits and
vegetables. He worked with a large number of fruits and vegetables during
his career in his endeavor to elucidate the physiological and biochemical
bases for quality maintenance. His papers have been milestones for others
to follow. Over time, he became increasingly interested in the less obvious
characteristics of flavor and nutritional quality. He realized that an improve-
ment in human diets through the consumption of more fruits and vegetables
would occur only when the appearance quality of fruits and vegetables was
matched by an increase in their flavor and nutritive quality. He became a
vocal proponent for coupling the traditional studies for improved appear-
ance quality with studies of flavor and nutritional quality changes during
harvest, storage, transport, and marketing. All this because he recognized
the importance of bringing to the end user a product that not only looks
great but also tastes wonderful and is at its optimal nutritive value.

D ed ic ati o n

Since beginning as a graduate student, Adel maintained a deep and

abiding interest in studying postharvest physiology and ensuring that fresh
fruits and vegetables are available to consumers in the best possible con-
dition. While a research assistant and as a technician, he was associated
with the late Dr. E.C. Maxie in pioneering studies on the effects of gamma
irradiation on the storability of fresh produce. His research was focused on
the responses of fruits to stress caused by O2 and CO2 during postharvest
handling. It involved the postharvest physiology of fruits, including mode
of action of oxygen and carbon dioxide on respiratory metabolism, ethylene
biosynthesis and action, and phenolic metabolism of fruits. Dr. Kader and
his coworkers developed new models and indices to predict fruit tolerance
for combinations of factors in controlled environments. They developed a
database for modified atmosphere packaging of fresh produce and iden-
tified optimum O2, CO2, and ethylene concentrations for storage of stone
fruits, Asian pear, kiwifruit, strawberry, and other commodities.
Professor Kader’s distinguishing characteristics included an amaz-
ing capacity to assimilate and organize information. He was universally rec-
ognized as among the most knowledgeable scientists in his field; however,
he was also widely known for freely sharing his knowledge and experience
with both the scientific community and the public. Intelligence, qualifica-
tion, organization, precision, and punctuality were the work tools that led
him to reach extraordinary scientific results. He also took the time to help
people and form lasting relationships. He was widely acknowledged for his
enthusiasm in teaching and in sharing his knowledge with others without
self-interests. At the age of 70, he organized many one-week short courses
and seminars with a few colleagues in Spain, Italy, Greece, India, Malaysia,
and UC Davis.
Few individuals, if any, can match the standards set by Professor
Kader during his tenure as a mentor. When it came to research integrity and
reaching for the highest standards possible, he practiced what he preached.
He demanded the highest performance from all colleagues, whether at UC
Davis or elsewhere. He took his responsibilities as a mentor very seriously.
One of Professor Kader’s most long-standing and sincerely held beliefs was
that students should be encouraged and given the means to attend scien-
tific meetings as a way of inspiring them in their developing careers. He
provided funding for countless students to travel to professional meetings
and advocated for allocating funds to support student travel, thus providing
the framework for the individual student to establish networking skills with
other scholars. Dr. Kader always had a gift handy in his bag: a T-shirt, a box
of chocolates, herb infusions, pens, a needle thermometer for fruit tempera-
ture, and sometimes the bag itself. Especially with young students, not only
did he dispense his knowledge through his lectures and seminars, but also
he spread “hard knowledge.” We all remember him recognizing the young-
est in the audience and presenting them with CDs, pen drives, or hard-copy

D ed ic ati o n

books. During his years at UC Davis, he had a constant stream of interna-

tional visitors. He fostered an interactive environment for students and vis-
iting scientists, helping students to further develop networking skills and
expanding their vision beyond the boundaries of UC Davis.
Professor Kader’s in-depth knowledge of postharvest was a challenge
in itself to anyone discussing topics of mutual interest. He challenged not
only his students and visiting scientists, but also the larger worldwide post-
harvest community, to question research results and resolve the seemingly
unsolvable questions that inevitably arise from research. He instilled a
desire to learn in his students. The positive aspect of this challenging envi-
ronment was the intellectual growth that occurred, because he nurtured
this growth by positive feedback, thoughtful comments, and critiques.
Adel Kader was a constant and inspiring role model. He was always
organized (everyone was in awe of his perfectly organized desk and his
ability to instantly find a journal reference in his files), always prepared,
and always ready to listen to anyone or extend a helping hand. From the
perspective of postharvest biologists, Dr. Kader’s signature achievement
has to be the development of the UC Davis Postharvest Technology Center.
From a loose affiliation of postharvest extension specialists, who published
sporadic issues of a postharvest bulletin, he developed what is widely recog-
nized as the world’s best source for postharvest information and education.
His vision established the annual postharvest short course, which is now
in its 35th year. The impact of this course, with more than 2500 alumni,
including students, researchers, teachers, regulators, and postharvest
practitioners from around the world, is incalculable. To ensure the center’s
continued vitality, he also developed, and was a tireless advocate for, the UC
Davis Postharvest Program Endowment Fund.
Professor Kader was born in Cairo, Egypt, in 1941. He earned his BSc
in horticulture from the Faculty of Agriculture at Ain Shams University in
Cairo in 1959. He started as a medical student, but at 14 years of age he
found he couldn’t stomach human organs and blood; thus, he transferred
to the Faculty of Agriculture. After obtaining his BS from Ain Shams, he
moved to UC Davis, where he earned his MSc in vegetable crops in 1962
and his PhD in plant physiology in 1966 at the age of 25. After earning his
doctorate, Professor Kader returned to Egypt, where he held the position
of assistant professor in the Faculty of Agriculture at Ain Shams University
from 1966 to 1971. While there, he engaged in teaching and research on
postharvest horticulture and coauthored a classic postharvest textbook in
Arabic. He then became a lecturer and consultant in the Kuwait Institute
for Scientific Research from 1971 to 1972, before returning to UC Davis in
1972, first as an assistant researcher and later as an assistant, associate,
and full professor until his retirement in 2007. He held the title of emeritus
professor until his death. During his tenure at UC Davis, he served as chair-
man of the Department of Pomology from 1986 to 1991, a member of the

D ed ic ati o n

campus academic planning council, and on numerous committees of the

UC system-wide Division of Agriculture and Natural Resources.
Dr. Kader received numerous awards. A listing of just a few of them
illustrates the breadth of his accomplishments. He was elected a fellow of
the American Society for Horticultural Sciences (ASHS) in 1986, later serv-
ing as president-elect in 1994–1995, president in 1995–1996, and chairman
of the board of directors in 1996–1997, as well as on the finance commit-
tee and as chair of the publications committee. Dr. Kader also received
awards for best research publications in 1978 and 1980 from the ASHS.
He was the chair of the program committee when UC Davis hosted the
22nd International Horticultural Congress in 1986. He attended every
controlled atmosphere conference, beginning with the second in 1977,
and convened the seventh edition of this conference in Davis in 1997. He
received from UC Davis the Outstanding Teaching Award in Extension
in 1989, the Award of Distinction from the College of Agricultural and
Environmental Sciences in 2000, the Alumni Citation of Excellence from
the Cal Aggie Alumni Association in 2000, and the Academic Senate’s
Distinguished Graduate Mentoring Award in 2003. He was also selected
as the Outstanding Horticulturist of 1997 by the Horticultural Research
Center at Laval University, Quebec, Canada. In April 2010, he received an
honorary doctorate degree from the University of Cartagena in Spain. In
2012, he was honored by the government of Malaysia for his outstanding
contributions to postharvest science, education, and extension.
As a member of several professional societies, Professor Kader served
on the editorial boards of Postharvest Biology and Technology, International
Journal of Postharvest Technology and Innovation, Postharvest News and
Information, and Tropical Science.
He cooperated with international organizations such as the Food and
Agriculture Organization (FAO) and United Nations Organization (UNO),
government programs such as the U.S. Agency for International Development
(USAID), and foundations such as the Gates Foundation. He worked with
many countries, including Saudi Arabia, Egypt, Syria, Iraq, India, Lebanon,
Mexico, Turkey, Morocco, Ghana, Sudan, Philippines, Thailand, Malaysia,
China, Chile, Jordan, and Kuwait. His laboratory and his home hosted a
continuing stream of visiting scientists from around the world. During his
university career, he trained 36 PhD students and more than 60 postdoc-
toral researchers who came from all over the world. Professor Kader served
for 18 years on the selected group of the Scientific Advisory Council of the
World Foods Logistics Organization, where he voluntarily supported the
industry all over the world.
Adel’s energy was also essential to the publication of the first and
subsequent editions of the companion text Postharvest Technology of
Horticultural Crops, third edition. More than 5700 copies in English have
been sold, and the text has also been translated into Spanish. When he

D ed ic ati o n

died, he was coordinating the writing of the fourth edition, his signature
book. He also served as author and editor of many publications, ­including
the popular Small-Scale Postharvest Handling Practices: A Manual for
Horticultural Crops, which he coauthored with Lisa Kitinoja and was trans-
lated into 11 additional languages. He was also associate editor of the book
Dates: Postharvest Science, Processing Technology and Health Benefits, which
was published after his death. Adel led the development of the Postharvest
Technology website (, which has become
the premier place to find postharvest information and receives several mil-
lion page views annually.
In teaching and research, Adel was a wonderful colleague. He and
his students were particularly focused on understanding the physiology,
biochemistry, and technology of controlled and modified atmosphere stor-
age of fruits and vegetables. He held the highest ethical, professional, and
research standards for both himself and others, which challenged everyone
with whom he worked to perform to their highest possible level. In addition
to hundreds of peer-reviewed papers and popular articles describing his
research, Adel, with photographer Don Edwards, also produced hundreds
of high-quality slides demonstrating his research findings. These slides are
still an essential component of many of the presentations made by members
of the UC Davis postharvest team. We would like to remember his intellec-
tual integrity and his way of discussing and interacting with colleagues and
students. He was happy to share his knowledge, he had no secrets, and he
believed research and information should be shared by and with everyone.
He was happy to give a bibliographic reference, photos, PowerPoint presen-
tation, publications, and so on.
Adel was convinced that improved postharvest practices would not
only raise the quality, taste, and nutrition of fruits and vegetables in the
United States, but also improve food supply and farmers’ incomes in the
developing world. From the start of his career, he was constantly involved
in development activities. As a key player in the Agriculture Development
Strategy (ADS) project, which sought to bring the expertise of U.S. hor-
ticulturists to Egypt, he made postharvest handling a central theme. The
subsequent flourishing of horticulture and horticultural exports from his
home country can, at least in part, be attributed to his efforts.
After retirement, Professor Kader maintained a very active interest in
postharvest programs worldwide and frequently participated in seminars at
UC Davis and many international meetings, chaired the California Citrus
Quality Council and the research advisory board of the Produce for Better
Health Foundation, and was board director of the Postharvest Education
Foundation. He continued to do some consulting to raise funds for the UC
Davis Postharvest Endowment. For the past several years, Adel served as a
key player in the Global Horticulture Assessment, which laid the foundation
for the development of the Horticultural Collaborative Research Support

D ed ic ati o n

Program (Horticulture CRSP), which USAID awarded to UC Davis. As an

advisor during the writing of the proposal, and as a member of its inter-
national advisory board, he made significant contributions to the success
and direction of the program. In nearly every project in which Adel was
involved, he took care to empower others working alongside him, and it is
due to this foresight that many of these projects were, and will continue to
be, completed to fruition, to the benefit of numerous others.
With the sudden death of Adel Kader on December 10, 2012, the
postharvest and horticultural development community mourned the loss
of a leader, teacher, mentor, colleague, and friend. His big heart, which
he shared so willingly with everyone, finally failed him while traveling
home from a postharvest conference in South Africa. In August 2014, at
a general assembly during the International Horticultural Congress in
Brisbane, Australia, Professor Kader was awarded an International Society
for Horticultural Sciences (ISHS) fellow posthumously, for his outstand-
ing contribution to horticultural science in general and postharvest science
and technology in particular during his long and distinguished career.
Professor Kader was instrumental in providing anyone who had the
privilege of working with him the necessary tools to feel at home in the
scientific community and become a contributor to the body of knowledge of
plant science. He also provided everyone in horticulture, and postharvest
biology in particular, something more important—the reinforcement of a
personal code of ethics that is crucial for being a member of the human
race. His high level of professional conduct, the care he took in mentoring
his students, and his humble approach to life are what enabled him to have
such an impact, not only on the world of postharvest biology, but also on the
world in general. Last but not least, it can be said that if postharvest is the
religion, Dr. Kader is the bible.

Elhadi M. Yahia, Mikal E. Saltveit, and Sunil Pareek

Series Preface

The ‘Innovations in Postharvest Technology’ book series provides

updated and comprehensive information on the innovations and emerging
technologies in postharvest and processing of horticultural commodities,
as well as postharvest physiology, biochemistry, ripening, and engineer-
ing aspects. The series includes books on ripening physiology, biochemis-
try, treatments to enhance shelf life, chilling injury, fresh cut and minimal
processing, postharvest pathology, and physiological disorders. The series
also includes books on postharvest biology and technology of tropical, sub-
tropical, and temperate fruits of global importance, as well as vegetables and
spices. Books also cover several aspects of general interest, such as supply
chain management, postharvest technology status in various regions of the
world, analytical techniques, biotechnology, engineering, nondestructive
quality evaluation, and health effects. The books are aimed at food scientists,
postharvest researchers and industries, and graduate- and postgraduate-
level students.


Horticultural crops have myriad uses in societies worldwide; not only are
fruits and vegetables critical components of the diet, but together with flow-
ers and ornamentals, they are pleasing aesthetic products that contribute
to human well-being. Edible products are a source of antioxidant vitamins
(A, C, and E), phenolics, carotenoids, phytonutrients, and dietary fiber that
are important health-promoting compounds in the human diet. The con-
tribution that these products make to human health has become increas-
ingly recognized and includes reduction of the incidences of degenerative
diseases as well as cardiovascular disease, hypertension, and cancers.
However, no amount of information about the “goodness” of any fruit or
vegetable is useful if it is not visually appealing or flavorful or does not meet
the quality expected by the consumer.
Access to horticultural products can vary greatly, with scarcity in
some areas and surplus in others. Estimates for losses of horticultural
crops vary widely, but it is interesting from survey research that losses of
fruits and vegetables are similar in the developing and developed worlds.
Yet, where those losses occur, between the “farm and fork,” can be mark-
edly different. In the developing world, losses are largely a result of fac-
tors such as lack of harvest, storage, and transport infrastructure and poor
marketing systems. In the developed world, losses are greater after harvest
because of stringent quality standards and factors such as excess produc-
tion, loss of product quality after harvest, and “plate waste” (unconsumed
food after purchase by consumers). Interestingly, the locavore movement,
or seeking locally grown food, often organic, in countries such as the United
States, can sometimes resemble the situation in developing countries, with

Fo re wo rd

small-scale and hobby farms lacking access to proper refrigeration and

appropriate marketing channels.
Selection for biological and physiological properties that ensure bet-
ter storage and transport capability of horticultural crops has often led
to products that are firm and slow to ripen, but with less sensory appeal,
such as relatively poor flavor and aroma. Also, the compromise between
quality and storage potential that exists for many fruits can result in their
harvest well before full-quality characteristics are attained in order to
maximize storage periods. Therefore, quality in the marketplace is often
at minimum acceptable levels. Together with selection of crops for yield
and uniformity of appearance, rather than consumer quality has resulted
in the paradox that consumers in developed countries have more choice
than any time in history, and yet dissatisfaction with food quality is high.
In part, this has led to the locavore movement because of the desire of con-
sumers to eat horticultural products that have not traveled long distances
and are perceived as fresher and healthier.
It is in this context that books such as Postharvest Ripening Physiology
of Crops provide an opportunity to summarize our understanding of the
complex interactions that result in ripening. Our ability to modify these
processes by breeding, whether traditionally or by manipulation of specific
genes by technology, requires ongoing research. In addition, understand-
ing of ripening processes has underpinned the development of existing
postharvest technologies and will be essential for development and imple-
mentation of newer technologies. Our understanding of ripening and
senescence processes in horticultural crops has progressed substantially
beyond the descriptive knowledge of the recent past. This progress has
been due to advances in omics technologies that are allowing identifica-
tion of genomic, proteomic, and metabolomic events that initiate and modu-
late these processes. The chapters in this book address these aspects, but
notably, morphological and physiological factors that affect ripening, and
thereby responses to postharvest environments, are also covered. Much
remains to be learned as the need to concurrently breed horticultural crops
with high-quality characteristics, while providing the ability to maintain
quality throughout storage, transport, and shelf life, and reduce waste, in
order to deliver high-quality crops with health benefits to the consumer,
will continue to increase.

Christopher B. Watkins

Cornell University, Ithaca, New York


Ripening is an important aspect of the production and shelf life of fresh pro-
duce. Timing and stage of ripening affect the buying behavior of produce,
shelf life, quality, and nutraceuticals. Ripening physiology is complicated
and affected simultaneously by many factors. Looking at these facts, this
book provides information on the postharvest physiology, biochemistry, and
molecular biology of ripening. Quality, physiology, and molecular biology
of flower senescence are also discussed. Detailed information on advances
in respiration measurement, stomatal relations in postharvest, and factors
controlling postharvest water loss has been provided. Lysophospholipids
research is gaining popularity and provides information on ripening and
extending shelf life of horticultural produce. A detailed account on posthar-
vest quality in relation to lysophospholipids is also included.
This book is a comprehensive interdisciplinary reference source for
the various aspects of fruit ripening and postharvest behavior. It focuses on
the postharvest physiology and ripening overview of fruits and vegetables.
Flower senescence is equally important for the postharvest horticulture
industry, and chapters are included on the postharvest quality of ornamen-
tal plants and molecular biology of flower senescence. The share of colored
fruit in the total fruit production is quite significant, with the largest con-
tributing several billion dollars annually. This has encouraged scientists
to study the pigmentation mechanism in fruits. Various developments that
have taken place in the last decade with respect to identifying and altering
the function of ripening-related genes are described in this book. Taking
clues from studies in grape as a model fruit, we review a few case studies,


and a detailed account of molecular regulation of fruit ripening, signal

transduction, and internal atmospheres in relation to fruit ripening is given.
Comparative proteomics is a useful tool to gain information on the
molecular events taking place during fruit maturation, in addition to find-
ing biotechnological strategies to improve horticultural traits, such as fruit
quality, shelf life, and yield. An overview of methods utilized in fruit pro-
teomics, as well as a global proteome and systems biology analysis of fruits
during ripening, is also presented. Potato is an important food crop of the
world; potato dormancy is associated with postharvest storage. The basics
of dormancy, its molecular and physiological basis, and methods to break
the dormancy are also discussed in detail in this book.
This book provides an overview of the most important metabolic path-
ways and genes that control volatile biosynthesis in model fruits, including
tropical, subtropical, and temperate fruits, with a special emphasis on fruit
ripening and the role of ethylene during this process. Also presented is
a brief description of the composition of volatiles in various fruit species
and a discussion of the influences of preharvest factors and postharvest
technologies on fruit aroma. A basis for product flavor, basic mechanisms
responsible for postharvest flavor change in fresh produce, and the poten-
tial impacts of various postharvest technologies on flavor are addressed.
I am sure that this book will serve as an important comprehensive
reference work for researchers, students, postharvest industries, and any-
one else who is involved in maintaining the postharvest quality of fresh

Sunil Pareek

Associate Professor (PHT)

Department of Agriculture & Environmental Sciences
National Institute of Food Technology Entrepreneurship and Management
(Deemed University under Ministry of Food Processing Industries)
Haryana, India


I gratefully acknowledge the special collaborations and suggestions made

by the scientists from the Maharana Pratap University of Agriculture and
Technology (MPUAT), Udaipur, India; National Institute of Food Technology
Entrepreneurship and Management (NIFTEM), Kundli, Haryana, India;
and Indian Council of Agricultural Research (ICAR), New Delhi, India.
Much appreciation is expressed to Dr. Ajit Kumar, vice chancellor, NIFTEM;
Dr. K.L. Chadha, former deputy director general (Horticulture Sciences),
ICAR; Dr. S.K. Malhotra, horticulture commissioner, Government of India;
Dr. H.P. Singh, former deputy director general (Horticulture Sciences),
ICAR; and Dr. S.L. Mehta, former deputy director general (Education),
ICAR, and vice chancellor, MPUAT, Dr. R. Paliwal, professor at Rajasthan
Agricultural Research Institute, Jaipur, India, and Dr. J.G. Varshney, head,
Department of Agriculture and Environmental Sciences, NIFTEM, Kundli,
India. The financial support by the ICAR under the National Agricultural
Research Project is deeply appreciated.
I am grateful to each of the authors for their participation, prompt-
ness, patience, and cooperation, as well as many others for their contribu-
tions, advice, and encouragement in the development of this book. I would
like to thank Ashley Weinstein and Stephen Zollo (CRC/Taylor & Francis)
for their support and encouragement in the preparation of the book ­proposal
and manuscript.
With my head stopped, I feel a paucity of words to express my
humble sense of regard to my parents, Mrs. Sushila and Dr. R.G. Pareek.
Finally, I think words are insufficient to express the feelings of my heart to

Ac k n o w l e d g m e n t s

acknowledge my better half, Dr. Shilpi, and son, Mr. Sabhya, who under-
went all sorts of hardships and sufferings to support my spirit and endeavor
at every step.


Dr. Sunil Pareek earned his PhD in horticulture

(PHT) from Rajasthan Agricultural University,
Bikaner, India. He joined Maharana Pratap University
of Agriculture and Technology (MPUAT), Udaipur,
India, in 2005 as an Assistant Professor (PHT) in the
Department of Horticulture, Rajasthan College of
Agriculture, MPUAT, Udaipur, India and worked there
up to August, 2015. Presently he is working as associ-
ate professor (PHT) in the Department of Agriculture
and Environmental Sciences, National Institute of
Food Technology Entrepreneurship and Management (NIFTEM), Kundli,
He is involved in teaching UG, PG, and PhD students, with a spe-
cial focus on postharvest physiology, technology, and processing of fruits.
He has guided several MSc and PhD students in PHT. He has executed
research projects supported by the Indian Council of Agriculture Research,
World Bank, Ministry of Tribal Development, and Rajasthan Mission on
Livelihood. Currently he is a co-consortium principal investigator (co-PI)
of the National Agricultural Innovation Project (NAIP) on underutilized
fruits, PI of the All India Coordinated Research Project on Tuber Crops, PI
of the Processed Products Scheme, and co-PI of the Integrated Farming
Systems Project.
Dr. Pareek has generated many technologies for the extension of the
shelf life of indigenous fruits and their processed products. Presently, he is
involved in a research program on applications of postharvest physiology of

Ed i t o r

fruits to maintain their quality during storage. He standardized and com-

mercialized the browning free pulp extraction and preservation technol-
ogy for sugar apple fruit. Dr. Pareek has published more than 40 papers,
40 ­presentations in national and international seminars and conferences,
6 books, 3 manuals, 6 technical bulletins, and 40 popular articles, and he
has several book chapters to his credit. He is the founder life member of
the Indian Society of Arid Horticulture and a member of several scientific
societies. He is on the reviewers’ panel of 12 international journals of repute
and also the editor, associate editor, or editorial board member of 11 inter-
national journals. He is the recipient of the University Outstanding Services
Award 2013, Young Scientist Award 2012, HS Mehta Young Scientist Award
2012, Fellow Award of the Confederation of Horticultural Associations of
India, and a few best poster awards.


Sasan Aliniaeifard Ajay Arora

Department of Horticulture Division of Plant Physiology
College of Abureyhan ICAR-Indian Agricultural Research
University of Tehran Institute
Tehran, Iran New Delhi, India

Domingos P.F. Almeida Jose G. Barbosa

Instituto Superior de Agronomia Department of Phytotechnology
Universidade de Lisboa Federal University of Viçosa
Lisbon, Portugal Viçosa, Minas Gerais, Brazil

Jane Ambuko Michael A. Campbell

Department of Plant Science and School of Science
Crop Protection Penn State Erie, The Behrend
College of Agriculture and College
Veterinary Services Erie, Pennsylvania
University of Nairobi
Nairobi, Kenya Bruno G. Defilippi
Instituto de Investigaciones
Fernanda F. Araujo Agropecuarias (CRI La Platina)
Department of Plant Physiology La Pintana, Santiago, Chile
Federal University of Viçosa
Viçosa, Minas Gerais, Brazil

Co ntributo rs

Sergio Tonetto de Freitas Kazuo Ichimura

Brazilian Agricultural Research NARO Institute of Floricultural
Corporation–Embrapa Science
Embrapa Tropical Semi-Arid National Agriculture and Food
Petrolina, Pernambuco, Brazil Research Organization
Fujimoto, Tsukuba, Japan
Cassandro Vidal Talamini do
Amarante Emrul Kayesh
Department of Agronomy Department of Horticulture
Santa Catarina State University Bangabandhu Sheikh Mujibur
Lages, Santa Catarina, Brazil Rahman Agricultural University
Gazipur, Bangladesh
Fernando L. Finger
Department of Phytotechnology Ola Lasekan
Federal University of Viçosa Faculty of Food Science and
Viçosa, Minas Gerais, Brazil Technology
University Putra Malaysia
Charles F. Forney Serdang, Malaysia
Atlantic Food and Horticulture
Research Centre Claudius Marondedze
Agriculture and Agri-Food Canada Department of Biochemistry
Kentville, Nova Scotia, Canada University of Cambridge
Cambridge, UK
Christoph Gehring
Division of Biological and Uulke van Meeteren
Environmental Sciences and Department of Plant Sciences
Engineering Wageningen University
King Abdullah University of Wageningen, the Netherlands
Science and Technology
Thuwal, Kingdom of Saudi Arabia Elizabeth J. Mitcham
Department of Plant Sciences
Orianne Gudenschwager University of California
Instituto de Investigaciones Davis, California
Agropecuarias (CRI La Platina)
La Pintana, Santiago, Chile Nora L. Olsen
Twin Falls Research and Extension
M. Mofazal Hossain Center
Department of Horticulture University of Idaho
Bangabandhu Sheikh Mujibur Twin Falls, Idaho
Rahman Agricultural University
Gazipur, Bangladesh

Co ntributo rs

Willis O. Owino Lingfei Shangguan

Department of Food Science and College of Horticulture
Technology Nanjing Agricultural University
Faculty of Agriculture Nanjing, China
Jomo Kenyatta University of
Agriculture and Technology Kenichi Shibuya
Nairobi, Kenya NARO Institute of Floricultural
Rakesh Pandey National Agriculture and Food
Division of Plant Physiology Research Organization
ICAR-Indian Agricultural Research Fujimoto, Tsukuba, Japan
New Delhi, India Tania P. Silva
Department of Phytotechnology
Federal University of Viçosa
Sunil Pareek
Viçosa, Minas Gerais, Brazil
Department of Horticulture
Rajasthan College of Agriculture
Jeffrey C. Suttle
Maharana Pratap University of
USDA-ARS Northern Crop Science
Agriculture and Technology
Udaipur, Rajastha, India
Fargo, North Dakota
Ludivine Thomas
Department of Agriculture &
Bioscience and Bioengineering
Environmental Sciences
Core Facility
National Institute of Food
King Abdullah University of
Technology Entrepreurship and
Science and Technology
Management (NIFTEM)
Thuwal, Kingdom of Saudi Arabia
Kundli, Sonepat, Haryana, India
Peter M.A. Toivonen
Vijay Paul Pacific Agri-Food Research Center
Division of Plant Physiology Agriculture and Agri-Food Canada
ICAR-Indian Agricultural Research Summerland, British Columbia,
Institute Canada
New Delhi, India
Christopher B. Watkins
Mikal E. Saltveit School of Integrative Plant
Mann Laboratory Science—Horticulture Section
Department of Plant Sciences Plant Science Building
University of California Cornell University
Davis, California Ithaca, New York

Co ntributo rs

Aloysius Wong Elhadi M. Yahia

Division of Biological and Human Nutrition Program
Environmental Sciences and Faculty of Natural Sciences
Engineering Autonomous University of
King Abdullah University of Queretaro
Science and Technology Juriquilla, Queretaro, Mexico
Thuwal, Kingdom of Saudi Arabia

Chapter 1

Ripening Physiology:
An Overview
Sunil Pareek 1,2
1Maharana Pratap University of Agriculture and
Technology, Udaipur, Rajasthan, India
2National Institute of Food Technology Entrepreneurship
and Management, Ministry of Food Processing
Industries, Kundli, Sonepat, Haryana

Abstract 2
1.1 Introduction 2
1.2  Climacteric Phenomenon 4
1.3 Physicochemical and Metabolic Changes 12
1.3.1  Color Changes 13
1.3.2  Sugar Changes 14
1.3.3  Organic Acid Changes 16
1.3.4  Flavor and Aroma Changes 18
1.3.5  Cell Wall and Textural Changes 22
1.3.6  Physiological Changes 28
1.4  Conclusions and Future Perspectives 33
References 33


Ripening is an important event in the fruit life cycle and from the consum-
ers’ point of view. Mature fruit undergo ripening, which is a coordinated
process typically involving many changes, such as pigmentation, flavor,
aroma, respiration, ethylene, texture, softening, sugar, and organic acid.
Physiologists typically classified fruits into climacteric and nonclimacteric
categories on the basis of their respiration peaks and internal ethylene
concentrations. This chapter reviews the climacteric and nonclimacteric
ripening of fruits and physiological and metabolic changes during ripen-
ing. In climacteric fruit, components responsible for the production of cli-
macteric ethylene have been identified. Less progress has been made on
nonclimacteric fruit; still, knowledge is poor to classify many fruits into
climacteric or nonclimacteric. A comprehensive climacteric classification
of fruits is given in this chapter. Textural changes and the role of enzymes
are also reviewed.

1.1 Introduction
Ripening is the composite of the processes that occur from the latter stages
of growth and development through the early stages of senescence and that
result in characteristic aesthetic and food quality, as evidenced by changes
in composition, color, texture, or other sensory attributes. Fruit ripening
is a complex process that undergoes dramatic changes mainly involving
flavor, color, and texture. It is a controlled process where the cellular com-
munication is important, as well as the influence of plant growth regulators
and environmental signals (Cruz-Hernandez and Paredes-Lopez, 2012).
Ripening could also be defined as a physiological process that is geneti-
cally programmed and comprises several physical, chemical, and biochemi-
cal changes that render fruit attractive and palatable (Lelievre et al., 1997;
Giovannoni, 2001).
One early view of ripening was that it is due to entirely to a series
of catabolic reactions associated with changes in membrane permeabil-
ity and a decrease in the structural integrity of the cell, resulting in the
release or activation of hydrolytic enzymes. This view is now clearly
untenable since genetic studies suggest that the expression of specific
genes is also required for ripening. It is supported by biochemical evi-
dence that shows there are changes in specific mRNAs and the de novo
synthesis of protein during ripening. Changes in gene expression are
both positive and negative. They involve genes in the nucleus and plas-
tids, and expression of at least some of them is confined to ripening fruit
tissues. Thus, fruit ripening is a highly controlled and programmed

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

developmental event, involving the coordination of a multitude of meta-

bolic changes (Seymour et al., 2002) and the activation and inactivation
of various genes (Clendennen and May, 1997; Medina-Suarez et al., 1997;
Drury et al., 1999; Itai et al., 2000), leading to various biochemical and
physiological changes within the tissue.
At ripening, fruits undergo many changes (Table 1.1), which, although
variable among species, generally include modification of cell wall ultra-
structure and texture, conversion of starch to sugars, alterations in pig-
ment biosynthesis and accumulation, and heightened levels of flavor and
aromatic volatiles (Brady, 1987).

Table 1.1  Changes Occurring in Fruit Ripening

Changes Events
Color Loss of chlorophyll
Dismantling of photosynthetic apparatus
Synthesis and accumulation of pigments
Texture Solubilization of pectin and cellulose
Starch degradation
Changes in protein content
Hydration of cell walls
Cell wall enzyme activity
Flavor and aroma Accumulation of sugars and organic acids
Production of volatiles
Alcohol ester synthesis
Control of pathways Increase in respiration
Ethylene synthesis
Changes in metabolism of starch and organic
Altered regulation of existing metabolic pathways
Gene expression Ripening-specific mRNA synthesis
Small and interference RNA appearance
Disappearance of mRNAs
Protein expression Synthesis of de novo ripening-specific proteins
Disappearance of proteins


1.2  Climacteric Phenomenon

Fruits can be broadly classified as climacteric or nonclimacteric, depend-
ing on whether or not a fruit exhibits a peak in respiration and ethylene
production during ripening. Climacteric fruits are characterized by a tran-
sient increase in ethylene synthesis and respiration at an early stage of
ripening. The peak of the ethylene production rate is proportional to the
peak respiration rate. An increased rate of respiration that occurs at an
early stage in ripening is termed climacteric, the term currently used to
describe the totality of events occurring during the ripening of climacteric
fruit. The study of the respiratory curve of climacteric fruits at a suitable
temperature shows a decreasing trend to the lowest value, termed the pre-
climacteric minimum, followed by a rise in respiration to the climacteric
peak and the subsequent postclimacteric decline in the rate of respiration
(Figure 1.1). During the ripening of climacteric fruits, the increase in ethyl-
ene and respiration is an early event (Figure 1.2). In many cases, occurring
prior to any changes in color or texture, this peak respiration corresponds
with eating ripeness, as has been observed in avocado, banana, cherimoya,
and mango. Fruit softening, color changes, development of taste and flavor,

Climacteric peak
Carbon dioxide production



Non-climacteric pattern


Figure 1.1  Climacteric pattern of respiration.

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Climacteric fruit

180 Breadfruit


ml O2 or CO2/Kg-Hr






20 Tomato

0 2 4 6 8 10 12 14 16 18
Time units

Figure 1.2  Respiration rate of some climacteric fruits.

and a number of other parameters of the ripening process are associated

with the climacteric cycle. Climacteric is therefore defined as a period in
the ontogeny of fruit during which a series of biochemical changes are
initiated by autocatalytic production of ethylene, making the change from
growth to senescence and involving an increase in respiration, leading to
ripening of the fruit (Payasi and Sanwal, 2005). Nonclimacteric fruit does


not show any increase in respiration and ethylene synthesis during ripen-
ing (Figure 1.3). In fact, nonclimacteric fruits show decline in their respira-
tion rate and ethylene production throughout the ripening process.
Climacteric events are mainly regulated by the gaseous phytohor-
mone ethylene, which is also involved in the decrease in flesh firmness
typical of many economically relevant crops, such as tomato and peach
(Prinsi et al., 2011). On the other hand, ripening of nonclimacteric fruits
such as pepper, citrus, and strawberry is ethylene independent, although
similar major visual, texture, flavor, and metabolic changes occur, as in cli-
macteric fruits. Many of the changes have been mainly characterized in
climacteric ripening fruits, whereas nonclimacteric fruit ripening is still
poorly understood. A list of climacteric and nonclimacteric fruit updated
from Biale and Young (1981), Kays (1991), and Watkins (2002) is shown
in Table  1.2. Interestingly, this physiological behavior is not linked to

Nonclimacteric fruit


ml O2 or CO2/Kg-Hr





0 1 2 3 4 5 6 7 8 9 10
Time units

Figure 1.3  Respiration rate of some nonclimacteric fruits.

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Table 1.2  Climacteric and Nonclimacteric Classification of Fruit

Common Name Scientific Name Reference
Acerola Malpighia emarginata DC. Carrington and King
Apple Malus pumila Mill. Biale (1960)
Apricot Prunus armeniaca L. Biale (1960)
Asian pear Pyrus serotina Rehder Downs et al. (1991)
Araza Eugenia stapitata Hernandez et al. (2007)
Atemoya Annona squamos x Brown et al. (1988)
Annona cherimola
Avocado Persia americana Mill. Biale (1960)
Bael Aegle marmelos (L.) Corr. Roy (1975)
Banana Musa spp. L. Biale (1960)
Biriba Rollinia deliciosa Safford Biale and Barcus (1970)
Bitter melon Momordica charantia L. Kays and Hayes (1978)
Blackberry Rubus spp. L. Walsh et al. (1983)
Blueberry, Vaccinium angustifolium Ismail and Kender (1969)
lowbush Ait.
Blueberry, Vaccinium corymbosum L. Ismail and Kender (1969)
Blueberry, Vaccinium ashei Reade Lipe (1978)
Breadfruit Artocarpus altilis Biale and Barcus (1970)
Caimito Pouteria caimito Malik et al. (2014)
Caja Spondias mombin L. Sampaio et al. (2007)
Camu-camu Myrciaria dubia Kunth Hernandez et al. (2009)
Canistel Pouteria campechiana Malik et al. (2014)
Cantaloupe Cucumis melo L. Lyons et al. (1962)
Cape gooseberry Physalis peruviana L. Trinchero et al. (1999)
Cherimoya Annona cherimola Mill. Biale (1960)
Chili plum Spondias purpurea var. Sampaio et al. (2007)
Corossol sauvage Rollinia orthopetala A. DC. Biale (1976)

(Continued )


Table 1.2  Climacteric and Nonclimacteric Classification of Fruit

Common Name Scientific Name Reference
Dabai Camarium odontophyllum Ding and Tee (2011)
Date palm Phoenix dactylifera L. Abbas and Ibrahim (1996)
Durian Durio zybethinus Murray Ketsa and Daengkanit
Feijoa Feijoa sellowiana O. Berg. Biale (1960)
Fig Ficus carica L. Marei and Crane (1971)
Giant granadilla Passiflora quadrangularis Malik et al. (2014)
Golden apple Spondias dulcis Forst. syn. Graham et al. (2004)
Spondias cytherea Sonn.
Golden berry Physalis peruviana L. Trinchero et al. (1999)
Guava Psidium guajava L. Akamine and Goo (1979)
Guava, ‘purple Psidium littorale var. Akamine and Goo (1979)
strawberry’ longipes (O. Berg.) Fosb.
Guava, Psidium rittorale Raddi Akamine and Goo (1979)
Guava, ‘yellow Psidium rittorale var. Akamine and Goo (1979)
strawberry’ littorale Fosb.
Honeydew melon Cucumis melo L. Inodorus Pratt and Goeschl (1968)
Jackfruit Artocarpus heterophyllus Selvaraj and Pal (1989)
Jujube, Chinese Ziziphus jujuba Mill. Kader et al. (1982)
Jujube, Indian Ziziphus mauritiana Lamk. Pareek and Yahia (2013)
Kiwifruit, Chinese Actinidia deliciosa (A. Pratt and Reid (1974)
gooseberry Chev) C.F. Liang et A.R.
Ferguson var. deliciosa
Lucuma Ponteria lucuma (Ruiz and Yahia (2004)
Pav.) Kuntze
Mammey apple Mammea americana L. Akamine and Goo (1978)
Mango, African Irvingia gabonensis Bailon Aina and Oladunjoye
& Irvingiaceae (1993)
Mango, common Mangifera indica L. Biale (1960)
Mangosteen Garcinia mangostana L. Paull and Ketsa (2004)
Monstera Monstera deliciosa Malik et al. (2014)
Nance Byrsonima crassifolia (L.) Velasquez de Klimo
Kunth (2001)
(Continued )

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Table 1.2  Climacteric and Nonclimacteric Classification of Fruit

Common Name Scientific Name Reference
Papaw Asimina triloba (L.) Dunal. Biale (1960)
Papaya Carica papaya L. Biale (1960)
Passion fruit, purple Passiflora edulis Sims Biale (1960)
Passion fruit, Passiflora edulis f. Malik et al. (2014)
yellow flavicarpa
Peach Prunus persica (L.) Batsch Biale (1960)
Pear, Chinese Pyrus breschneideri R. Tian et al. (1987)
Pear, European Pyrus communis L. Biale (1960)
Persimmon Diospyros kaki L. Reid (1975)
Plum Prunus americana Marsh. Biale (1960)
Raspberry Rubus idaeus L. Burdon and Sexton (1990)
Sapodilla Manilkara achras (Mill.) Malik et al. (2014)
Sapote Casimiroa edulis Llave Biale (1960)
Sapote, black Diospyros digyna Jacq. Yahia (2004)
Sapote, mamey Pouteria sapota Jacq. H.E. Yahia (2004)
Moore & Stearn
Sapote, white Casimiroa edulis Llave & Yahia (2004)
Saskatoon Amelanchier alnifolia Nutt. Rogiers et al. (1998)
Soursop Annona muricata L. Biale and Barcus (1970)
Sweetsop, sugar Annona squamosa L. Brown et al. (1988)
Tomato Lycopersicon esculentum Biale (1960)
Achachairu Garcinia humilis (Vahl) Durate (2011)
C.D. Adam
Asian pear Pyrus serotina Rehder Downs et al. (1991)
Blackberry Rubus spp. L. Perkins-Veazie et al.
Cacao Theobrama cacao L. Biale and Barcus (1970)
Cactus pear Opuntia amyclaea Tenore Lakshminarayana and
Estrella (1978)
Carambola Averrhoa carambola L. Lam and Wan (1983)
Cashew Annacardium occidentale L. Biale and Barcus (1970)
(Continued )


Table 1.2  Climacteric and Nonclimacteric Classification of Fruit

Common Name Scientific Name Reference
Cherry, sour Prunus cerasus L. Blanpied (1972)
Cherry, Surinam Eugenia uniflora Malik et al. (2014)
Cherry, sweet Prunus avium L. Biale (1960)
Cranberry Vaccinium macrocarpon Ait. Kader (2002)
Cucumber Cucumis sativus L. Biale (1960)
Grape Vitis vinifera L. Biale (1960)
Grapefruit Citrus paradise Macf. Biale (1960)
Gumichama Eugenia brasiliensis Malik et al. (2014)
Indian gooseberry Emblica officinalis Gaertn. Pareek and Kitinoja
Jaboticaba Myrciaria cauliflora (Mart.) Mota et al. (2002)
O. Berg
Java plum Syzygium cuminii (L.) Skills Akamine and Goo (1979)
Lemon Citrus jambhiri Lush. Biale (1960)
Litchi (Lychee) Litchi chinensis Sonn. Akamine and Goo (1979)
Longan Dimocarpus longan Lour. Zhao et al. (2005)
Loquat Eriobotrya japonica Lindl. Pareek et al. (2014)
Malay apple Syzygium malaccense Malik et al. (2014)
Mandarin Citrus reticulata Blanco Reid (1975)
Mountain apple Syzygium malaccense (L.) Akamine and Goo (1979)
Merrill & Perry
Olive Olea europeae L. Maxie et al. (1960)
Orange Citrus sinensis (L.) Osb. Biale (1960)
Pepper Capsicum annuum L. Saltveit (1977)
Pineapple Ananas comosus (L.) Merr. Biale (1960)
Pitanga Eugenia uniflora L. Santos et al. (2006)
Pitaya Hylocereus spp. Nerd et al. (1999)
Pitaya, yellow Selenicereus megalanthus Nerd and Mizrahi (1999)
Scum. Ex Vaupel
Pomegranate Punica granatum L. Ben-Arie et al. (1984)
Pummelo Citrus grandis Malik et al. (2014)
Rambutan Nephelium lappaceum L. Mendoza et al. (1972)
Raspberry Rubus idaeus L. Perkins-Veazie and
Nonneeke (1992)
Red bayberry Myrica rubra Sieb. Zucc. Joyce and Li (2003)
(Yang mei)
(Continued )

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Table 1.2  Climacteric and Nonclimacteric Classification of Fruit

Common Name Scientific Name Reference
Rose apple Syzygium jambos (L.) Alston Akamine and Goo (1979)
Star apple Chysophyllum cainito L. Pratt and Mendoza (1980)
Strawberry Fragaria x ananassa Duch. Biale (1960)
Strawberry, wild Fragaria vesca L. Nam et al. (1999)
Surinam cherry Eugenia euniflora L. Akamine and Goo (1979)
Tamarind Tamarindus indica Yahia (2004)
Tomato (nor-, rin-, Lycopersicon esculentum Thompson et al. (1999)
cnr-) Mill.
Tree tomato, Cyphomandra betacea Pratt and Reid (1976)
Tamarillo (Cav.) Sendtu
Watermelon Citrullus lanatus (Thumb.) Elkashif et al. (1989)
Wax apple Syzygium samarangense Akamine and Gao (1979)
(Blume) Merr. and L.M.

taxonomic groups. Species belonging to the same family, such as tomato

and pepper (Solanaceae), display a distinct response to ethylene. Thus,
tomato is a climacteric fruit while pepper is not (Palma et al., 2011). With
other fruit, such as kiwifruit, a hybrid ripening pattern is seen, with most
of the ripening changes occurring in the absence of any detectable rise in
ethylene and CO2 production; a climacteric response occurs only toward
the end of ripening. Exposure to exogenous ethylene promotes ripening of
kiwifruit, but if exposure to ethylene is insufficient or fruit are too imma-
ture, then removal of ethylene results in nonclimacteric behavior.
White (2002), while reviewing fruit development and ripening, pro-
vides evidence from biochemical and genetic studies that both ethylene-
dependent and ethylene-independent regulatory cascades control the
development of tomato fruit. Hence, although the nonripening tomato
mutants ripening-inhibitor (RIN ) and non-ripening (NOR) do not produce
autocatalytic ethylene or ripen in the presence of exogenous ethylene, they
do display signs of ethylene sensitivity and ethylene-inducible expression
of several genes. Thus, it is likely that RIN and NOR participate in ethylene-
independent regulatory cascades during the early stages of fruit ripening
(White, 2002). Jim Giovannoni and colleagues have isolated the RIN and
NOR genes by positional cloning strategies (Moore et al., 2002; Vrebalov
et al., 2002). LeMADS-RIN encodes a member of the MADS-box family of
transcription factors. Homologues of LeMADS-RIN are expressed during
the ripening of other fruit, including strawberry, which might indicate a


common (ethylene-independent) function in the ripening of both climac-

teric and nonclimacteric fruit.
Several papers describe the role of ethylene in the ripening of climac-
teric fruit (Alexander and Grierson, 2002; Klee, 2002; Moore et al., 2002;
Payasi and Sanwal, 2009; Bapat et al., 2010; Bouzayen et al., 2010; Pech
et al., 2012, 2013). These concentrate on the elucidation of biochemical and
genetic signaling cascades that impact the development and ripening of
tomato fruit. The isolation of transcription factors NOR and LeMADS-RIN,
which participate in ethylene-independent signaling in tomato, and the dis-
covery that a homologue of the RIN gene is expressed in nonclimacteric
fruit have suggested that the common regulatory cascades may operate
in all fruits. This knowledge could enable generic strategies to manipulate
the ripening of any fruit (White, 2002). Such work has been complemented
by transcriptional profiling during the development and ripening of both
climacteric and nonclimacteric fruit (Aharoni and O’Connell, 2002; Moore
et al., 2002; Seymour et al., 2002), which may disclose more common regu-
latory elements.
Nevertheless, the timing of the climacteric syndrome in nonclimac-
teric fruit is not related to the ripening period per se. In grape, it occurs
at veraison (Chervin et al., 2004), and in citrus, in young immature fruit
(Katz et al., 2004). In strawberries, ethylene production starts to increase
once the fruit reaches the red ripe stage, but not before (Lannetta et al.,
2006). It is now considered that some aspects of the ripening of noncli-
macteric fruit are regulated by ethylene, and in climacteric fruit, some
ripening pathways are independent of ethylene action (Pech et al., 2013).
Differential cross talk between ethylene and other phytohormones prob-
ably operates in each type of fruit (refer to Chapter 12 for details) (Pech
et al., 2008; Paul et al., 2012).

1.3 Physicochemical and Metabolic Changes

Ripening changes involve a multiplicity of biochemical, metabolic, and
molecular changes that affect the cell compartments (Table 1.1): they have
been shown to be related to alterations in specific enzymes or complete
pathways. Color changes, for example, are due to alterations in the chloro-
phyll and pigment content of the plastids (Carrilo-Lopez et al., 2003; Zhang
et al., 2006). Softening is brought about by alteration in cell wall metabolism
and is due to a partial solubilization of pectin and cellulose; starch degra-
dation may contribute to a change in texture (Carrilo-Lopez et al., 2002).
Alterations in the metabolism of organic acids and the generation of volatile
compounds that produce aroma are common. It is clear that the various
changes associated with ripening take place in different parts of the cell
and are highly coordinated.

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

1.3.1  Color Changes

Changes in fruit color typically involve the destruction of chlorophyll to
reveal other pigments already present and may also involve the synthesis
of additional pigments (Table 1.3). More than 25 years ago, Von Loesecke
revealed that the yellowing of banana peels during ripening is essentially
an unmasking of preexisting carotenoids as chlorophyll is broken down,
although more recent work has shown that subtle changes do occur in the
various carotenoids present. In tomato, carotenoid synthesis accompa-
nies the loss of chlorophyll, while in fruits that turn purple, blue, or black,
anthocyanins are synthesized as chlorophyll is depleted. In tomato, where
this process has been well studied and represents the transformation of
the chloroplast into a chromoplast, carotenoid synthesis has been shown
to be light dependent (Giovannoni, 2001). In some fruits, ripening may also
involve a change in the fruit surface waxes, altering the bloom or shininess
of the fruit (Carrington, 2011). Table 1.4 provides the carotenoid concentra-
tions in various fruits.
Conversion of chloroplast-rich photosynthetic fruit to a chromoplast
and nutrient-rich, nonphotosynthetic fruit is essential for the development

Table 1.3  Carotenoid Derivatives Synthesized during Ripening in Different

Carotenoid Derivative Fruits
Lycopene Tomato, watermelon, guava, papaya, grapefruit
β-Carotene Cantaloupe melon, mango, pumpkin, apricot
Zeaxanthin Orange pepper, citrus fruit, persimmon
Capsanthin Red pepper
Capsorubin Red pepper

Table 1.4  Range of Carotenoid Concentration in Some Fruits

Carotenoid Concentration
Fruit (mg 100 g−1) Reference
Apricot 0.1–4 Kurz et al. (2008)
Tomato 10.5–27.8 Ilahy et al. (2011)
Citrus species 0.02–5 Fanciullino et al. (2008)
Papaya 1.5–3 De Souza et al. (2008)
Loquat 0.1–2 Zhou et al. (2007)
Pepper 0.2–20 Topuz and Ozdemir (2007)
Watermelon 3–7 Perkins-Veazie et al. (2001)


of storage compartments (i.e., chromoplasts) with increased capacity to

accumulate large quantities of the lipophilic carotenoids (for more details,
refer to Egea et al., 2010). During the chloroplast-to-chromoplast transition,
specific carotenoid biosynthesis genes are expressed. The first step in carot-
enoid biosynthesis corresponds to the condensation of two geranylgera-
nyl diphosphate molecules into phytoene, which is catalyzed by phytoene
synthase (PHY ). In tomato fruit, two PHY genes are expressed. Phytoene
synthase 1 (PHY1) is highly expressed in ripening fruit and is responsible
for the formation of chromoplastic carotenoids, while phytoene synthase 2
(PHY2), which is responsible for the formation of chloroplastic carotenoids,
is expressed exclusively in green tissues, and therefore makes no contri-
bution to carotenoid biosynthesis in ripening fruit (Fraser et al., 1999). In
tomato, the isoprenoid biosynthetic pathway gives rise to carotenoids, such
as β-carotene and lycopene, gibberellins, quinines, and sterols. This path-
way has been well researched since its manipulation impacts not only the
organoleptic qualities of fruit, but also their contribution to human health.
Other red and purple pigments of the type seen in grapes and boy-
senberries are anthocyanins, which are products of the phenylpropanoid
pathway. Anthocyanin pigments are water soluble, synthesized in the cyto-
sol, and localized in the vacuole. Their basic ring structure can be modified
by hydroxylation, methylation, or glycosylation, and their specific color is
modified by pH, metal ions, and co-pigments to produce the subtlety of col-
ors seen in nature. Anthocyanin content varies significantly in a range of
fruits (Table 1.5).
The accumulation of anthocyanins is regulated by transcription fac-
tors of two classes (R2R3 MYB and basic helix loop helix), regulatory pro-
teins that coordinate gene expression of the whole phenylpropanoid pathway.
In fruit, this regulation system has been well characterized in grape and
apple. In white berry grapes, VvMYBA2 is inactivated by mutations in the
coding region and VvMYBA1 has a retrotransposon in the promoter and is
not transcribed (Kobayashi et al., 2004; Walker et al., 2007). In apple fruit,
a mini-­satellite repeat structure in the promoter region of the MYB10 gene
upregulated the expression of this regulatory gene, which increased the level
of anthocyanin throughout the plant, producing a fruit with striking red color
throughout the flesh (Espley et al., 2009) (for details, refer to Chapter 9).

1.3.2  Sugar Changes

During fruit ripening, sugar levels within fruit tend to increase (Whiting,
1970), due to either increased sugar importation from the plant or the mobi-
lization of starch reserves within the fruit, depending on the fruit type and
whether it is ripened on or off the plant. The sugar or sugar alcohol, deliv-
ered to the fruit, is converted to starch (e.g., mango, banana, kiwifruit),

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Table 1.5  Anthocyanin Content in Common Fruits

Fruit Anthocyanin (mg 100 g−1 FW)
Acai 53.6
Acerola 22.6
Apple, Fuji 0.7
Apple, Gala 1.1
Apple, Golden 0.0
Apple, Granny Smith 0.0
Apple, Red Delicious 3.8
Avocado 0.3
Blackberry 90.6
Blueberry 141.0
Cherry 27.7
Cranberry 85.5
Currant, black 154.8
Currant, red 75.0
Currant, white 0.0
Elderberry 485.3
Grape, Concord 65.6
Grape, red 44.0
Grape, green 0.0
Kiwifruit 0.0
Melon 0.0
Nectarine and peach 1.8
Pear 12.2
Pineapple 0.0
Plum, red 6.98
Plum, black 39.7
Plum, yellow 0.3
Raspberry 40.9
Strawberry 23.8
Watermelon 0.0
Note: FW, Fresh Weight.

stored as reducing sugar (e.g., tomato, strawberry), or stored as sucrose

(e.g., wild tomato, watermelon, grape), or may even be converted to ­lipids
(e.g., olive). Accumulation of sucrose, glucose, and fructose in fruits
such as melon, watermelon (Brown and Summers, 1985), strawberry


Table 1.6  Starch Mobilizing Enzymes and Their Response in Fruits

Enzymes Mode of Action
α-Amylase Hydrolyzes the α (1–4) linkages of amylase at random to
produce a mixture of glucose and maltose
β-Amylase Attacks the penultimate linkage and releases only maltose
Starch Hydrolyzes the terminal α (1–4) linkage to give glucose-1-
phosphorylase phosphate, which can be converted to glucose-6-
phosphate by the action of glucose phosphate mutase
α-1,6-Glucosidase Attacks α-1,6-glucose linkages of amylopectin
Source: Data from Garcia, E., and Lajola, A.M., Journal of Food Science 53,
1181–1186, 1988; Tucker, G.A., Biochemistry of Fruit Ripening,
1–52, 1993.

(Fait et al., 2008), and peach (Lo Bianco and Rieger, 2002) is evident dur-
ing ripening. The main sugar at the fruit ripening stage depends on the
plant species. Glucose is the major sugar in table grape, whereas fructose
is the predominant sugar in berries, mango, and citrus species. Those
fruits that accumulate fructose and glucose show very low concentrations
of sucrose. However, apricot, plum, nectarine, and peach have sucrose as
the main sugar, which is accumulated during stage 3 as a result of a rise
in the activity of sucrose synthase (Morandi et al., 2008). The proportions
of fructose, glucose, and sucrose are important in the perception of taste
since fructose is 80% sweeter than sucrose, whereas glucose is only 60%
sweeter than sucrose (Yamaguchi et al., 1970). A regular increase in the
level of sugars in pear throughout the period of fruit development has been
observed due to translocation of photosynthates from leaves to the young
fruit, which are partly used for the synthesis of pectic substances and other
cell wall materials and partly converted to the usual storage product, the
starch. With the advancement of maturity, the accumulated starch is hydro-
lyzed into sugars, which is known as a characteristic event for fruit ripening
(Hulme, 1958). The starch-degrading enzymes in fruits and their mode of
starch mobilization are given in Table 1.6. Further breakdown of sucrose
into glucose and fructose is probably mediated by the action of invertase.
Low levels of invertase activity have been linked to the sucrose accumula-
tion trait in tomatoes (Yelle et al., 1991).

1.3.3  Organic Acid Changes

The peculiar taste and flavor of fruit is dependent on the type of organic
acid, predominant organic acid, and ratio between the organic acid and
sugar. The predominant organic acid in ripe fruit varies among species.

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

As quoted by Etienne et al. (2013), “Malic acid is dominant in apple

(Yamaki, 1984), loquat (Chen et al. 2009) and pear (Lu et al. 2011), whereas
citric acid is dominant in citrus fruits (Yamaki, 1989). In many fruit spe-
cies, differences in total acidity or in the balance of organic acids among
cultivars are also observed, for example in loquat (Yang et al. 2011), peach
(Etienne et al. 2002), pear (Lu et al. 2011), citrus (Albertini et al. 2006),
pineapple (Saradhuldhat and Paull, 2007), apricot (Gurrieri et al. 2001) and
banana (Bugaud et al. 2011).” Loss of acidity implies decarboxylation of
carboxylates, which can occur through the conversion of tricarboxylates
into dicarboxylates, but also through decarboxylation of the dicarboxylates
malate and oxaloacetate (OA A), leading to the degradation of organic acids
(Figure 1.4). Decarboxylation of OA A and malate allows the production of

NAD-ME acetylCoA CoA
acetylCoA citrate citrate
OAA CS citrate OAA
ACO glyoxylate
cycle isocitrate isocitrate
malate NAD-MDH
malate malate ICL
TCA Cycle
fumarate MS
isocitrate glyoxylate
succinate CoA NADP-IDH
citrate ATP-CL acetylCoA
GABA shunt
ACO acetylCoA
NADP-ME isocitrate
Pyruvate malate
NADP-IDH flavonoids/
2-oxoglutarate isoprenoids
PEPCK glutamate
glucose PEP OAA

Figure 1.4  Citrate and malate metabolic pathways in fruit mesocarp cells.
The probable direction of reversible reactions is indicated by the large
arrow. ACO, aconitase; ATP-CL, ATP-citrate lyase; CS, citrate synthase; ICL,
isocitrate lyase; MS, malate synthase; NAD-MDH, NAD-malate dehydro-
genase; NAD-ME, NAD-malic enzyme; NAD-IDH, NAD-isocitrate dehy-
drogenase; NADP-ME, NADP-malic enzyme; NADP-IDH, NADP-isocitrate
dehydrogenase; PDH, pyruvate dehydrogenase; PEPC, phosphoenolpyru-
vate carboxylase; PEPCK, phosphoenolpyruvate carboxykinase; PPDK, pyru-
vate orthophosphate dikinase. (Adapted from Etienne, A. et al., Journal of
Experimental Botany, doi:10.1093/jxb/ert0352013.)


phosphoenolpyruvate (PEP) and is linked to the activation of gluconeogen-

esis (Sweetman et al., 2009). Gluconeogenesis is a m ­ etabolic pathway that
results in the generation of glucose from PEP. It occurs mostly during fruit
ripening when sugars accumulate rapidly (Sweetman et al., 2009).
The downward trend in the levels of organic acids with the onset of
fruit ripening has been reported by a number of workers. The total organic
acid (malic + citric + quinic) decreased in peach with the ripening of fruits,
coinciding with decreasing titratable acidity (Wang et al., 1993). Being a
major respiratory substrate, organic acid (predominating) is metabolized
to a greater extent than the others and may fall by 50% during the life of the
fruit. Malic acid, as a major substrate of respiration, has been suggested,
and thus accounts for the respiratory quotient of 1.1 or higher, which is
typical of pome fruit (Fidler and North, 1967). The decline in the content
of organic acids during fruit ripening might be the result of an increase in
membrane permeability, which allows acids to be stored in the respiring
cells (Kliewer, 1971), as well as formation of salts of malic acid, reduction in
the amounts of acid translocated from the leaves, reduced ability of fruits
to synthesize organic acids with fruit maturity (Hardy, 1968), translocation
into sugars (Hulme, 1970), and dilution effect due to the increase in volume
of fruit.

1.3.4  Flavor and Aroma Changes

Flavor is the most important factor determining whether consumers will
repurchase a particular fruit. Two main factors determine a fruit’s char-
acteristic flavor: the correct sugar–acid balance and the production of
aroma volatile compounds. These volatile compounds can include a mix-
ture of volatile acids, aldehydes, alcohols, esters, terpenoids, and aromatics
(Table 1.7).
Flavor changes deal with the generation of volatiles to alter aroma,
the generation of sugars from stored starch, the interconversion of sugars,
a decline in acidity through altered organic acid metabolism, and a reduc-
tion in astringency through lower levels of tannins and phenolics (Kays,
1991). Even in a fruit as relatively nonaromatic as breadfruit, some 40
distinct volatile compounds are produced during ripening (Iwaoka et al.,
1994). Typically, only a handful of these volatiles, termed character impact
compounds, are the source of the distinctive odors associated with this
fruit. The second aspect of flavor relates to carbohydrate and organic acid
metabolism, and mango provides a good example. As it ripens, starch is
broken down, with glucose being converted to fructose and sucrose, while
at the same time various organic acids, chief among these succinate, are
depleted (Lazan et al., 1993). Some fruits have high levels of soluble phe-
nolics when immature. These result in astringency; during ripening, these

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Table 1.7  Aroma Substances in Fruits and Description of Their Odor

Aroma Substance Odor Description
(E)-2-Hexenal ‘Hayward’ kiwifruit
1-Methyl ethyl butyrate Apple
2-Hexenal Green leaf, green banana
2-Methyl butyl acetate Apple
2,6-Nonadienal Cucumber
Acetaldehyde Pungent, penetrating
Acetone Sweet, pungent
Allyl isothiocynate Raw cabbage
Butyl acetate Fruity
Butyl butyrate Fruity, pear
Citral Lemon
Dimethyl disulfide Cooked cabbage
Dimethyl disulfide Onion, cabbage
Ethyl acetate Etherlike, pineapple, anise
Ethyl butanoate ‘Hort16A’ kiwifruit
Ethyl butyrate Fruity, pineapple
Ethyl hexanoate Fruity
Eugenol Ripe banana
Furaneol Strawberry
Heptanone Banana
Hexanal Cut grass
Hexenal Sweet, almond, green
Hexyl acetate Fruity, apricot
Isopentanol Overripe banana
Linalool Fruity, floral, citrus
Methoxypyrazine Grassy
Methyl anthranilate Foxy
Methyl butyrate Apple
Methyl hexanoate Etherlike, pineapple
Monoterpene limonene Lime
Nootakatone Grapefruit
Raspberry ketone Raspberry
Valencene Orange
α-Farnesene Apple
β-Damascenone Richness
β-Lanon-trans Warm, woody, balsamic, rose


are polymerized, and astringency is lost, as these tannins are no longer

capable of interacting with taste receptors (Kays, 1991).
Watson et al. (2002) noted that many volatile and nonvolatile com-
pounds give rise to the flavor of strawberry fruit. The volatile compounds
give the fruit its distinctive flavor, whereas the nonvolatile compounds, such
as sugars and organic acids, are responsible for the fruit’s sweetness and
tartness. In tomato, several sensory attributes have been proposed to char-
acterize aroma, such as fruity, green, grassy, earthy, musty, floral, candy,
citrus, grapefruit, and pharmaceutical aromas (Causse et al., 2001; Baldwin
et al., 2004; Sinesio et al., 2010). More than 400 aroma volatiles have been
identified in tomato fruit, among which about 30 seem to be important for
tomato aroma (Baldwin et al., 2000, 2004). In apple and strawberry, more
than 300 volatile compounds have been identified (Dixon and Hewett,
2000), for which 20 volatiles are considered the aroma fingerprint of straw-
berry (Ulrich et al., 1997).
Amyl esters give bananas their distinctive flavor and aroma, and
butyl esters give them a fruity flavor and aroma; however, other esters and
aldehydes, alcohols, and ketones have been associated with flavor, and their
production rates can increase during ripening (Tressl and Jennings, 1972).
McCarthy et al. (1963) also claimed that amyl esters gave bananas their
distinctive flavor and aroma, and butyl esters gave them a fruity flavor and
aroma. Volatile compounds of several banana cultivars have been widely
studied by many authors: ‘FLHORBAN 920’ and ‘Grand Naine’ (Bugaud
et al., 2009); ‘Gran Enana’, a subgroup of the ‘Cavendish’ originating from
Central and South America (Vermeir et al., 2009); various cultivars grown
on Madeira Island (Nogueira et al., 2003); banana fruits (Musa sapientum L.
var. ‘Cavendish’) from Honduras and their aqueous essences (Jordan et al.,
2001); free and glycosidically bound volatile compounds of the ‘Valery’ and
‘Pequeña Enana’ cultivars (Pérez et al., 1997); banana fruits (Musa caven-
dishii L.) of the ‘Gran Enana’ and ‘Enana’ cultivars from the Canary Islands
and the ‘Enana’ cultivar from Colombia (Cano et al., 1997); Philippine
bananas (‘Del Monte’, ‘Cavendish’); Taiwanese bananas (cv. ‘Sen-nin’); and
‘Delicious’ bananas (hybrid between ‘Philippine’ and ‘Taiwanese’) (Shiota,
1993). Generally esters such as butyl acetate, isoamyl acetate, ethyl acetate,
butyl butanoate, and isoamyl isobutanoate are responsible for the charac-
teristic aroma of fresh banana and constitute the major class of compounds
present in banana’s volatile profile (Salmon et al., 1996). The typical aroma
of banana is characterized by the presence of a wide range of volatile metab-
olites—with different volatilities and concentrations that can vary among
the different cultivars—as the initial work of Cano and collaborators with
Spanish and Columbian ‘Enana’ cultivars showed (Cano et al., 1997). They
identified the compounds and the ratios of the peak area to the internal stan-
dard area corresponding to the gas chromatography–mass spectrometry
(GC-MS) of the purge-and-trap analysis of three banana cultivars (Spanish

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

‘Enana’, Spanish ‘Gran Enana’, Latin American ‘Enana’). Quantifiable

­differences among the flavors of the banana cultivars were found. Spanish
‘Enana’ fruit was found to be the richest in flavor compounds. McCarthy
et al. (1963) classified the various components of banana aroma. A banana-
like flavor was assigned to the amyl and isoamyl esters of acetic, propionic,
and butyric acid, whereas the alcohols and carbonyls gave odors described
as green, woody, or musty. In their study, the Latin American banana flavor
displayed the presence of ethanol, 1-butanol, and hexanal, but these com-
pounds were not found in any of the Spanish banana varieties. Furthermore,
only Spanish bananas exhibited the presence of hexyl butanoate, which is
related to a banana-like flavor.
Pontes et al. (2012) evaluated the effect of the cultivar (‘Dwarf
Cavendish’, ‘Prata’, ‘Maçã’, ‘Ouro’, and ‘Platano’) on the volatile profile
determined by dynamic headspace solid-phase microextraction (dHS-
SPME) combined with one-dimensional gas chromatography–mass spec-
trometry (1D-GC-MS). This approach allowed the definition of a volatile
metabolite profile to each banana variety and can be used as pertinent
criteria of differentiation. A total of 68 volatile organic metabolites were
tentatively identified and used to profile the volatile composition in differ-
ent banana cultivars, thus emphasizing the sensitivity and applicability of
SPME for establishment of the volatile metabolomic pattern of plant sec-
ondary metabolites. Ethyl esters were found to comprise the largest chemi-
cal class, accounting for 80.9%, 86.5%, 51.2%, 90.1%, and 6.1% of total peak
area for ‘Dwarf Cavendish’, ‘Prata’, ‘Ouro’, ‘Maçã’, and ‘Platano’ volatile frac-
tions, respectively. de Vasconcelos Facundo et al. (2012) determined the
volatile differences in two cultivars of banana under cold storage conditions.
Cold storage more strongly affects the ‘Nanicão’ than the ‘Prata’ cultivar.
Esters such as 2-pentanol acetate, 3-methyl-1-butanol acetate, 2-methylpro-
pyl butanoate, 3-methyl butyl butanoate, 2-methylpropyl 3-methyl butano-
ate, and butyl butanoate were drastically reduced in the cold group of the
‘Nanicão’ cultivar.
A comparative study of volatile components of 9 litchi cultivars (10
samples) from southern China was carried out through GC-MS com-
bined with headspace SPME (Wu et al., 2009). A total of 69 volatiles were
detected, of which 43 were identified and another 53 were tentatively
identified: 35  terpenoids, 27 alcohols, 10 aromatic compounds, 9 alde-
hydes, 8  esters, 4  ketones, 2 sulfurs, and 1 organic acid. Seventeen com-
mon volatiles included in all the samples were linalool, cis-rose oxide,
R-terpineol, β-citronellol, geraniol, p-cymene, ethanol, 3-methyl-3-buten-
1-ol, 3-methyl-2-buten-1-ol, 1-hexanol, (E)-2-hexen-1-ol, 2-ethyl-1-hexanol,
1-octen-3-ol, 1-octanol, ethyl acetate, p,R-dimethylstyrene, and 3-tert-butyl-
4-hydroxyanisole. In these cultivars, ‘Guangxi Huaizhi’ contained at most
67 volatiles, and ‘Jizhuili’ contained at least 36 volatiles. Alcohols were the
predominant volatile components in various cultivars, representing 35.1%


(‘Guangdong Huaizhi’) to 81.6% (‘Jizhuili’) of the main fraction. Although

the volatile composition and concentration varied between these cultivars,
the components with the highest odor activity values (OAVs) in most culti-
vars were still cis-rose oxide, trans-rose oxide, 1-octen-3-ol, and geraniol.
Two Huaizhi samples from two producing areas exhibited similar volatile
profiles and were significantly different from other cultivars according to
cluster analysis performed on amounts of major volatile components (Wu
et  al., 2009). Fourteen volatile compounds, namely, three alcohols, one
ester, three monoterpenes, and seven sesquiterpenes, were detected at har-
vest in cultivar Mauritius in South Africa. Cultivar McLean’s Red showed
19 volatile compounds, namely, 4 alcohols, 3 monoterpenes, 2 oxides, and
10 sesquiterpenes. In cultivar Mauritius, alcohols represented 50% of the
main fraction, followed by sesquiterpenes (23%), monoterpenes (21%), an
unknown component (6%), and the ester (0.7%). No ester was detected in
cultivar McLean’s Red (Sivakumar et al., 2008). Among the three alco-
hols, citrinelol and geraniol predominated in the aroma profile of cultivar
Mauritius, conferring a characteristic “floral, rose, citrus and fruity aroma”
to the fruit (Chyau et al., 2003). Limonene, rose oxide, citronellol, and gera-
niol were detected at relatively low levels in cultivar Mauritius, although
rose oxide was not present in the latter. Zingiberene was the predominant
sesquiterpene, and terpinolene the most abundant monoterpene in cultivar
Mauritius (Sivakumar et al., 2008).

1.3.5  Cell Wall and Textural Changes

Recently several reviews (Vicente et al., 2007d; Goulao and Oliveira, 2008;
Li et al., 2010; Cruz-Hernandez and Paredes-Lopez, 2012) have been pub-
lished on the role of cell wall–degrading enzymes and textural changes.
Fruit softening and related textural alterations during ripening are mainly
consequences of progressive depolymerization and solubilization of cell wall
components and loss of cell structure. Both enzymatic and nonenzymatic
factors contribute to softening (Brummell and Harpster, 2001; Dumville
and Fry, 2003; Brummell, 2006). Moreover, different fruits soften at dif-
ferent rates and by varying degrees due to the inherent composition and
nature of their cell wall polysaccharides and other cell wall structural com-
ponents (Table 1.8) (Tucker and Grierson, 1987; Brummell, 2006). Fruit
such as banana, mango, kiwifruit, avocado, and papaya undergo dramatic
softening; fruit such as apple, grape, and citrus do not exhibit such drastic
changes. Different rates of softening can also be illustrated by varied dura-
tions required for loss of firmness for different fruit types; for example,
comparative studies showed that fruit softening at room temperature may
take 3 days for both tomato and banana, 4.5 days for mango, 30 days for
carambola, and 24 days for ‘Kampuchea’ guava (Ali et al., 2004).

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Table 1.8  Common Polymers of Primary Cell Wall and Their Structures
Polymer Molecular Structure
Cellulose (1→4)β-D-Glucan chains held together with
hydrogen bonding, forming very long crystalline
Cross section contains 36 glucan chains
Glucomannan Backbone contains regions of (1→4)β-D-glucan and
(1→4)β-D-mannan in nearly similar amounts
Occasionally terminal; has a side chain of single
units of α-D-galactose
Glucuronoarabinoxylan Backbone of (1→4)β-D-xylan
Side chains of single unit of nonreducing terminal
α-L-arabinose and α-D-glucoronic acid
Homogalacturonan Made of long chains of (1→4)α-D-galacturonic acid
Initially highly methyl esterified
Rhamnogalacturonan I Made of alternating α-D-rhamnose and α-D-
(RG-I) galacturonic acid residues; long side chains of
either unbranched (1→4)β-D-galactan or branched
α-L-arabinans or type I arabinogalactans attached
to the rhamnose residues
Rhamnogalacturonan II Backbone made of (1→4)α-D-galacturonic acid like
(RG-II) homogalacturohan; complex side chains of
different types of neutral sugar
A minor cell wall component
RG-II monomers can domimerize together as boron
diesters and may affect the cell wall porosity
Structural proteins Four different types including expansin; some are
heavily glycosylated
Xyloglucan Backbone similar to cellulose, i.e., (1→4)β-D-glucan
Regular substitution on three out of four consecutive
glucose residue with α-D-xylose
Xylose occasionally extended with β-D-galactosyl-α-L-
fucose or α-L-arabinose in some species
The reducing end of unsubstituted glucose residues
is susceptible to cleavage by Trichoderma endo-
(1→4)β-D-glucanase (EGases) producing similar
amounts of heptasaccharide (G1c4.Xy13) and
nonasaccharide (G1c4.Xy13.Gal.Fuc) xyloglucan
subunit oligosaccharides


Fruit cell walls consists of complex networks of polysaccharides and

proteins, wherein the primary cell wall contains, on average, about 35% pec-
tin, 25% cellulose, 20% hemicellulose, and 10% structural protein, depending
on the fruit species and developmental stage (Brownleader et al., 1999).
Major classes of cell wall polysaccharides are modified to varying levels
during fruit ripening, and a scheme depicting softening and those cell wall
modifications occurring in ripening has been proposed for a melting-flesh
peach by Toivonen and Brummell (2008). In general, cell wall modification
and depolymerization during fruit ripening are variable among different
fruits (Brummell, 2006; Toivonen and Brummell, 2008). For example, pec-
tin depolymerization has been reported as the major cause of loss of firm-
ness in raspberry and boysenberry (Vicente et al., 2007a, 2007c), whereas
only slight pectin depolymerization is detected in ripening strawberry
(Huber, 1984), banana (Wade et al., 1992), and blueberry (Vicente et al.,
2007b). Major cell wall enzymes assayed in different fruits during ripening
are given in Table 1.9.

Table 1.9  Cell Wall Enzymes Assayed in Fruit Ripening

Fruits Enzymes
Climacteric Fruits
Apple β-Galactosidase, glucanase, polygalacturonase, pectin
methyl esterase (PME)
Avocado Glucanase, polygalacturonase
Banana Glucanase, polygalacturonase, PME
Kiwi Polygalacturonase, xylanase (xyloglucan
Mango β-Galactosidase, PME
Melon β-Galactosidase, glucanase, polygalacturonase, PME
Papaya β-Galactosidase, glucanase, polygalacturonase, PME
Peach β-Galactosidase, glucanase, polygalacturonase, PME
Pear β-Galactosidase, β-glucosidase, polygalacturonase
Pepper Glucanase
Tomato Glucanase, polygalacturonase, PME, xilanase
Nonclimacteric Fruits
Grape α-Galactosidase, β-galactosidase, PME
Orange α-Galactosidase, β-galactosidase, glucanase,
α-glucosidase, β-glucosidase, PME
Prickly pear β-Galactosidase, glucanase, polygalacturonase, PME
Strawberry Glucanase, PME

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Softening in kiwifruit occurs over a period of weeks and can be

divided into a number of phases (Figure 1.5). Modifications of the cell wall
play an important part in determining fruit texture and ripening charac-
teristics. Chemical analyses of cell wall components show some consistent
changes during the early stages of ripening. These include

1. Solubilization of pectin (but without further degradation)

2. The cell wall swelling and showing an increased affinity for water
(becoming more hydrophilic)
3. Loss of galactose from pectins (especially of a galactan that is
tightly associated with the cellulose microfibrils)
4. Deesterification of some pectins

These changes continue once kiwifruit have begun rapidly softening to

ripeness (phase 2 in Figure 1.5). Phase 2 softening is associated with a
further increase in pectin solubilization, loss of galactan and arabinan side
chains from pectic polymers, and more cell wall swelling. As softening pro-
gresses into phase 3, two more important changes begin, both of which
appear to be regulated by ethylene:

5. Depolymerization (a reduction in size) of the hemicellulosic poly-

saccharide xyloglucan, which is associated with a reduction in cell
wall strength
6. Depolymerization of pectin, which is associated with dissolution of
the middle lamella and reduced intercellular adhesion.

These six changes have been observed in a wide range of fruit types,
although the extent and relative timing vary somewhat between species.
Such observations indicate that pectin solubilization and cell wall swelling
are important events in the control of softening in kiwifruit and probably
most other species with melting texture.
Graham Seymour and colleagues at HRI–Wellesbourne are investi-
gating the biochemistry of fruit texture in order to improve the palatabil-
ity and shelf life of produce (Marin-Rodriguez et al., 2002; Seymour et al.,
2002). Marin-Rodriguez et al. (2002) reviewed the role of pectate lyases in
fruit softening. Pectate lyases (PELs) catalyze the Ca 2+ -dependent cleavage
of deesterified pectin, which is a major component in the primary cell walls
of many higher plants. Initially, it was thought that these enzymes were
produced solely by plant pathogens to macerate plant tissues. As a result of
both plant genome sequencing and EST programs, however, it has become
clear that these enzymes are encoded by large gene families in plants and
expressed throughout the plant, including ripening fruit. Tomato, straw-
berry, grape, and banana fruits all express PELs, where they may play a
significant role in fruit softening. Other enzymes involved in modifying


Respond to exogenous ethylene Autocatalytic ethylene production

Starch degradation

Pectin solubilization
Fruit firmness (%)

60 Phase 1
Soluble pectin depolymerization
galactose loss

Aroma production
respiratory climacteric
Phase 2 pectin depolymerization
20 rapid softening
Loss of middle lamella
Phase 3
eating window Phase 4

Figure 1.5  Schematic representation of postharvest ripening in kiwifruit,

showing the timing of key physiological events. At harvest, fruit do not pro-
duce ethylene but are highly sensitive to exogenous ethylene. Softening is
initiated (phase 1) and becomes rapid (phase 2). Relatively late in softening,
compared with other fruit species, endogenous autocatalytic ethylene pro-
duction begins, aroma volatiles are produced, and fruit become soft enough
to eat (phase 3). If fruit progress to the overripe stage (phase 4), they become
unacceptably soft and exhibit off-flavor notes. (Reproduced from Atkinson
R.G. et al., Journal of Experimental Botany 62, 3821–3835, 2011.)

cell wall properties include pectin esterases (PEs) and polygalacturonases

(PGs). These enzymatic activities (Table 1.10) are similarly encoded by
multigene families, in which at least one member shows ripening-specific
Several studies have been carried out to investigate the role of cell
wall degradation-related enzymes and the expression of respective genes
during softening of the grape berries (Nunan et al., 2001; Ishimaru and
Kobayashi, 2002; Waters et al., 2005). Chervin et al. (2008) showed that
low doses of ethylene application increased the berry diameter at the incep-
tion of the ripening stage. The study verified previous studies that showed
correlation between berry ripening and the accumulation of various

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Table 1.10  Pectin-Modifying and -Degrading Enzymes in Fruits

Substrate Enzymes Products
Pectin Pectin methyl esterase Pectic acid + methanol
Endopolymethyl Methyl oligogalacturonides
galacturonase α-(1,2)-linked L-Rha-α-(1,4)-
linked D-Gal
Endopectinlyases Unsaturated
Hairy pectin Rhamnogalacturonan Pectin + acetic acid
acetyl esterase
Pectin acetyl esterase Pectin + acetic acid
Smooth pectins Lyases Oligogalacturonides
Protopectin Protopectinase Pectin
Pectic acid Endo-PG Oligogalacturonides
Exo-PG Monogalacturonides
Endopectate lyases Oligogalacturonides
Exopectate lyases Unsaturated
Trigalacturonic Oligogalacturonide Monogalacturonides
acid hydrolase
∆4:5 ∆4:5 unsaturated Unsaturated
(galacturonide) n oligogalacturonide monogalacturonide +
hydrolase galacturonides (n − 1)
Unsaturated Oligogalacturonide lyases Unsaturated
digalacturonate monogalacturonides
Arabinans α-L-Arabinofuranosidase α-L-Arabinose
(1,5)-α-Arabinans Endoarabinanase Arabinose and higher
Galactans β-D-Galactanase β-D-Galactose

transcripts of PG, PME, cellulose, and expansion. Some of the enzymes

may be involved in the increase of berry expansion by ethylene.
Manenoi et al. (2007) determined endixylanase gene expression, pro-
tein amount, and activity in three papaya cultivars that differ in softening
pattern and in one cultivar where softening was modified by the ethylene
receptor inhibitor 1-MCP. Differential expression of gene and enzyme activ-
ity was noticed in different cultivars and one treated with 1-MCP. The xyla-
nase gene could be utilized to control the growth and abscission.


1.3.6  Physiological Changes

On-tree and off-tree ripening physiology is dependent on respiration and eth-
ylene responses and concentration in fruit. As discussed earlier, fruits can be
classified into climacteric and nonclimacteric categories, depending on respi-
ration peaks and response to exogenous ethylene. Therefore, it is important to
study the respiration and ethylene behavior during ripening and postharvest
in fruits. Here respiration rates, ethylene concentrations, and their changes
during growth and ripening in various fruits are discussed (Table 1.11). Other
aspects related to physiological changes during on-tree ripening and posthar-
vest ripening are discussed throughout the book in various chapters.

Table 1.11  Respiration and Ethylene Production Rates Measured at 20°C in

Several Climacteric and Nonclimacteric Fruits
Respiration Rates Ethylene Production
Fruit (mg CO2 kg−1 h−1) (µlC2H4 kg−1 h−1)
Apple 20–31 25–2500
Apricot 40 < 0.01 (0°C)
Araza 1283 nd
Asian pear 25
Atemoya 250 200 (20°C)
Avocado 190 28.9–74.2
Banana 280 0.05–2.1
Blackberry 115 0.1–2.0
Blueberry 70 0.5–10.0
Breadfruit 480 (25°C) 1.2
Carambola 65 < 3.0
Cherimoya 300 200
Cherry, sweet 65 < 0.01 (0°C)
Cranberry 16 0.6 (5°C)
Dragon fruit 105 < 0.1
Durian 265 40
Fig 50 0.6 (0°C)
Gooseberry 81 nd
Grape, American 33 < 0.1
Grape, muscadine 51 < 0.1
Grape, table 27 < 0.1
Grapefruit < 10 (15°C) < 0.1

(Continued )

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Table 1.11  Respiration and Ethylene Production Rates Measured at 20°C in

Several Climacteric and Nonclimacteric Fruits
Respiration Rates Ethylene Production
Fruit (mg CO2 kg−1 h−1) (µlC2H4 kg−1 h−1)
Guava 74 10
Honeydew melon 30 Very low
Kiwifruit 19 75
Lemon 24 0.11–0.17
Lime nd 0.30–1.96
Litchi 60 Very low
Longan 42 Very low
Loquat 80 Very low
Mamey apple 35 (25°C) 400.0 (27°C)
Mandarin 25 < 0.1
Mango 58 0.04–3.0
Mangosteen 21 (25°C) 0.03
Nectarine 87 5.0 (0°C)
Netted melon 55.0 55.0
Orange 28 < 0.1
Papaya 80 8.0
Passion fruit 262 280.0
Peach 87 0.9–20.7
Pepper 34 < 0.2
Persimmon 22 < 0.5
Pineapple 24 0.16–0.40
Plum 20 0.14–0.23
Pomegranate 24 < 0.1
Prickly pear 32 0.2
Rambutan nd Very low
Raspberry 125 12.0
Sapodilla 16 (25°C) 3.7
Sapote nd >100
Star apple 38 0.1
Strawberry 150 < 0.1
Tomato 35 3.6–29.8
Watermelon 21 < 1.0
Wax apple 10 Very little
Note: nd, Not detected.


The respiration rate of avocado fruit is relatively high compared

to that of many other fruits: about 20–50 mg CO2 kg−1 h−1 at 5°C, 50–160
mg CO2 kg−1 h−1 at 10°C, and 80–300 mg CO2 kg−1 h−1 at 20°C (Kader and
Arpaia, 2001). Rates of ethylene production are generally low for unripe
avocados, < 0.1 µl kg−1 h−1 at 20°C, but increase rapidly after harvest up to
> 100 µl kg−1 h−1 at 20°C when fully ripe. Thus, the implications for ethylene
in avocados depend on the physiological state of the fruit.
Carambola exhibits nonclimacteric ripening behavior. Carambola
respiration was fairly constant at 20°C, < 20 ml CO2 kg−1 h−1 at 20°C
(Warren, 2009). It increased slightly while turning from yellow to orange,
likely owing to senescence of the fruit. Ethylene production was very low
at 25°C, about 0.4 µl kg−1 h−1. In an another study, fruit at the full-ripe (one-
quarter orange) stage had higher respiration and ethylene production rates
(Oslund and Davenport, 1983).
The respiration rate of dates is very low, < 5 mg CO2 kg−1 h−1 at 20°C
at the khalal stage and < 1 mg CO2 kg−1 h−1 at the rutab and tamr stages
(Yahia, 2004). Cured ‘Deglet Noor’ dates with 20%–22% moisture produced
0.4 mg CO2 kg−1 h−1 at 24°C and 2 mg CO2 kg−1 h−1 when the moisture con-
tent increased to 27% (Rygg, 1975). The rate of CO2 production is high ini-
tially, but declines steadily as the fruit advances in maturity, reaching its
lowest level as the fruit enters the stage of physiological maturity and then
increases to reach a peak as fruit ripens (Abbas and Ibrahim, 1996). Dates
produce very low concentrations of ethylene: < 0.5 µl kg−1 h−1 for the kha-
lal stage and < 0.1 µl kg−1 h−1 for the rutab and tamr stages kept at 20°C
(Yahia, 2004). Ethylene production in ‘Hallawi’ dates was not detected until
91 days after pollination; it increased to reach a peak within 15 days and
then declined rapidly (Abbas and Ibrahim, 1996).
Durian has a climacteric respiration characteristic with a peak rate
of around 200–250 mg CO2 kg−1 h−1 at 25°C; ethylene production ranges
between 0.3 and 1.4 µl C2H4 kg−1 h−1 during the preclimacteric stage and 8
and 12 µl C2H4 kg−1h−1 at its peak (Siriphanich et al., 1994). Internal atmo-
spheric composition changes with the ripening stage. CO2 increased from
1%–4% at harvest to 4%–14% when ripened at 22°C, while ethylene increases
from 0.5–1 µl L −1 to 3–7 µl L −1 (Tongdee et al., 1990).
Grape is a nonclimacteric fruit with a relatively low respiration rate
(3–4 ml CO2 kg−1 h−1 at 5°C). At 5°C, grapes produce 50% less heat of respi-
ration than other nonclimacteric fruits. Comparison of the cluster compo-
nents indicates that the respiration rate of the rachis alone is approximately
15 times higher than that of the berry at 4°C (Gardea et al., 1994). Grapes
produce very small amounts of ethylene during berry development. There
is peak of ethylene production at bloom, followed by a decreasing concen-
tration until harvest (Weaver and Singh, 1978).
During guava fruit ripening, ethylene production and respiration
rates of fruit harvested at the full-size, pale-green stage were higher than

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

those of fruit harvested at the small- or medium-size, dark-green stage

(Brown and Wills, 1983). Fruit harvested at advanced maturity reach their
respiratory climacteric stage in 4–6 days, accompanied by rapid changes
in skin color and flesh firmness. Ethylene production rates increase dur-
ing fruit ripening and reach a peak that may or may not coincide with the
respiratory peak. Guava cultivars such as ‘Allahabad Safeda’, ‘Apple Color’,
and ‘Hisar Safeda’ produce higher amounts of ethylene than ‘Lucknow-49’
(Singh and Pal, 2008).
Jackfruit respiration behavior follows a typical climacteric pattern
with three distinct phases of respiration (preclimacteric, climacteric, and
postclimacteric stages). Precut immature fruit slices showed a basal respi-
ration rate of 160 mg CO2 kg−1 h−1 fruit, and in the climacteric phase, the
peak of respiration was 245 mg CO2 kg−1 h−1 fruit (Selvaraj and Pal, 1989).
The climacteric peak was attained after 3 days of ripening. After 8 days of
ripening, the respiration rate had dropped to a residual level of 60–70 mg
CO2 kg−1 h−1 fruit.
Climacteric behavior in Chinese jujube is cultivar dependent. Some
of the cultivars show a climacteric nature of ripening, while few have
a nonclimacteric ripening pattern. Shen et al. (2004) reported that the
respiration rate and ethylene production of ‘Dongzao’ jujube at 4°C and
20°C each exhibited increasing tendencies with extension of storage time,
but that no apparent peaks of respiration or ethylene production were
detected. This suggested that ‘Dongzao’ jujube fruit is nonclimacteric.
In contrast, Jiang and Sheng (2003) found that the rates of respiration
and ethylene production of ‘Jins’ jujube during storage initially increased
and then reached respective peaks before finally decreasing rapidly; they
were further promoted when the fruit was exposed to ethylene. These
observations suggest that ‘Jins’ cultivar is climacteric. Indian jujube has a
high respiration rate and a climacteric respiration pattern (Pareek et al.,
2009; Pareek and Yahia, 2013). The rate of respiration of mature green
Indian jujube fruit of both Ziziphus mauritiana and Ziziphus spina-christi
is low but increases as the fruit matures, reaching a peak value when it
enters the ripening phase and then declining rapidly (Abbas, 1997). Abbas
and Fandi (2002) found that the respiration rate was high 3 weeks after
anthesis, but then declined steadily until 12 weeks after anthesis, when it
reached 14.2 mg CO2 kg−1 h−1. The rate then began to increase, first slowly
and then rapidly to a maximum value of 100.3 mg CO2 kg−1 h−1 as the fruit
entered the ripening phase. The rate of CO2 production declined rapidly
as the fruit became overripe. This pattern of respiratory change is charac-
teristic of a climacteric fruit.
Litchi fruit is nonclimacteric with relatively low levels of ethylene
production after harvest. The fruit does not ripen after harvest, and eth-
ylene production remains constant at 1°C–3°C storage temperatures for
30 days (Chen et al., 1968). According to Jiang et al. (1986), a decline in


the rate of respiration was observed during fruit development. In cultivar

‘Huaizi’, the respiration decreased directly after harvest, and thereafter
an increase was observed.
There is a controversial assessment concerning ethylene production
versus fruit maturity in loquat. Some authors found ethylene production
and an increase of respiration rate prior to color break (Gariglio et al., 2002;
Amoros et al., 2003), permitting Amoros et al. (2003) to define loquat as
a climacteric fruit. Hirai (1980) observed a marked increase in respira-
tion during fruit maturation, and the observation may have contributed to
the reclassification of loquat as a climacteric fruit by some authors, which
makes the classification of loquat controversial (Hasegawa et al., 2010).
However, others reported that although there is a well-defined peak of
ethylene production, it is insufficient to define loquat as a climacteric fruit
because there is not a concomitant increase in respiration rate, which often
appears ­irregular (Hamauzu et al., 1999; Ding et al., 1998; Kader, 2002;
Gonzalez et al., 2004). Therefore, there is no conclusive evidence to define
loquat fruit as a climacteric fruit, since (1) there is no starch accumulation at
stage 2 of fruit development (Gariglio et al., 2008); (2) there is no CO2 –C2H4
production relationship (Reig et al., 2007); (3) there is no general response
to e­ thephon to promote fruit ripening (Reig et al., 2007); (4)  there is no
confirmed evidence of an increase in hemicellulase, cellulose, and pectin
disrupting enzyme activities linked to ethylene production; and (5) there
is evidence of a nonclimacteric fruit postharvest behavior (Kader, 2002).
It is a well-established fact that mango exhibited a climacteric
pattern of respiration and an increase in ethylene production during
­r ipening. Respiration is very high after fruit set and then declines and
is maintained at a low rate until fruit ripening begins (Brecht and Yahia,
2009). Respiration patterns and ripening behavior vary among cultivars,
with different climatic conditions and growing locations. The respira-
tory peak in ‘Alphonso’ mangoes harvested when mature green occurs
5 days after harvest, and the fruit ripen within 7 or 8 days. ‘Piari’ man-
goes ripen on day 9 (Krishnamurthy and Subramanyam, 1970). The rise
in climacteric respiration in ‘Dashehari’, ‘Amrapali’, and ‘Rataul’ mangoes
coincides with the highest level of sucrose and PG activity in ripening
fruit (Kalra and Tandon, 1983). Mango fruit ripening is accompanied by
increased ethylene production, which coordinates the ripening process.
Mango expresses an autocatalytic increase in ethylene production during
ripening (Mattoo and Modi, 1969). Ethylene production starts before full
ripeness is reached (Cua and Lizada, 1990). Ethylene production in unripe
mango fruit is very low < 0.1 μl kg−1 h−1) (Burdon et al., 1996). Ethylene
production decreases as the fruit matures, is then undetectable for a
time, and reappears upon initiation of ripening (Akamine and Goo, 1979).
‘Kent’ and ‘Haden’ mango fruit have internal ethylene concentrations of
0.00  μl L −1 during the preclimacteric phase, increasing to 0.08 μl L −1 at

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

the initiation of the climacteric stage and up to 3.0 μl L −1 at the c­ limacteric

peak. Burg and Burg (1962) reported that ethylene production rises when
or before CO2 production rises in ripening mangoes, whereas Biale and
Young (1981) included mangoes among fruits in which ethylene rises
after CO2 production rises.
In papaya, the peaks in rates of respiration and ethylene production
coincide with full skin color development (Manenoi et al., 2007). Rates of
respiration and ethylene production began to rise after 2 days at 22°C in
‘Rainbow’ papayas were harvested at the color-break stage and reached
their maximum after 10 and 8 days of harvesting, respectively (Manenoi
et al., 2007). The number of days to reach respiratory and ethylene climac-
teric peaks and fruit ripening during postharvest also decreases with the
advancement of fruit maturity at harvest. For example, Golden’ papayas
harvested at different stages, viz. mature green, up to 15% yellow, up to 25%
yellow, and up to 50% yellow, reached the edible-ripe stage after 7, 6, 4, and
3 days at 23°C, respectively (Bron and Jacomino, 2006).

1.4  Conclusions and Future Perspectives

Remarkable progress has been made in understanding fruit ripening and
associated changes. Fruit flavor, aroma, softening, color, and sugar–acid
blend are important qualitative characteristics that determine the purchas-
ing of ripened fruit. Delaying ripening and maintaining quality throughout
the supply chain is the main objective of fruit physiologists and postharvest
technologists. Omics technology (proteomics, transcriptomics, genomics,
metabolomics) can be applied in this area. It has already provided a large
amount of information on metabolomic, proteomic, and transcriptomic
changes occurring during fruit ripening. However, very little has been
studied on systems biology for unraveling regulatory networks during fruit
ripening. A systems biology approach is needed for understanding climac-
teric and nonclimacteric fruit ripening. Epigenetics is another novel area
for fruit ripening study in the modern era.

Abbas, M.F. 1997. Jujube. In Mitra, S.K. (ed.), Postharvest Physiology
and Storage of Tropical and Subtropical Fruits. CAB International,
Wallinhgford, Oxfordshire, UK. pp. 405–415.
Abbas, M.F., and Fandi, B.S. 2002. Respiration rate, ethylene production
and biochemical changes during fruit development and maturation of
jujube (Ziziphus mauritiana Lamk). Journal of the Science of Food and
Agriculture 82, 472–476.


Abbas, M.F., and Ibrahim, M.A. 1996. The role of ethylene in the regulation
of fruit ripening in the Hillawi date palm (Phoenix dactylifera). Journal of
the Science of Food and Agriculture 72, 306–308.
Aharoni, A., and O’Connell, A.P. 2002. Gene expression analysis of strawberry
achene and receptacle maturation using DNA microarrays. Journal of
Experimental Botany 53, 2073–2087.
Aina, J.O., and Oladunjoye, O.O. 1993. Respiration, pectolytic activity and tex-
tural changes in ripening African mango (Irvingia gabonensis) fruits.
Journal of the Science of Food and Agriculture 63, 451–454.
Akamine, E.T., and Goo, T. 1978. Respiration and ethylene production in mam-
mee apple (Mammea americana L.). Journal of the American Society for
Horticultural Science 103, 308–310.
Akamine, E.T., and Goo, T. 1979. Respiration and ethylene production in fruits
of species and cultivars of Psidium and species of Eugenia. Journal of the
American Society for Horticultural Science 104, 632–635.
Albertini, M.V., Carcouet, E., Pailly, O., Gambotti, C., Luro, F., and Berti, L.
2006. Changes in organic acids and sugars during early stages of devel-
opment of acidic and acidless citrus fruit. Journal of Agricultural and
Food Chemistry 54, 8335–8339.
Alexander, L., and Grierson, D. 2002. Ethylene biosynthesis and action in
tomato: A model for climacteric fruit ripening. Journal of Experimental
Botany 53, 2039–2055.
Ali, Z.M., Chin, L.H., and Lazan, H.A. 2004. A comparative study on wall
degrading enzymes, pectin modifications and softening during ripening
of selected tropical fruits. Plant Science 167, 317–327.
Amoros, A., Zapata, P., Pretel, M.T., Botella, M.A., and Serrano, M. 2003.
Physico-chemical and physiological changes during fruit development
and ripening of five loquat (Eriobotrya japonica Lindl.) cultivars. Food
Science and Technology International 9, 43–51.
Atkinson, R.G., Gunaseelan, K., Wang, M.Y., Luo, L., Wang, T., Norling,
C.L., Johnston, S.L., Maddumage, R., Schroder, R., and Schaffer, R.J.
2011. Dissecting the role of climacteric ethylene in kiwifruit (Actinidia
­chinensis) ripening using a 1-aminocyclopropane-1-carboxylic acid
­oxidase knockdown line. Journal of Experimental Botany, 62, 3821–3835.
Baldwin, E.A., Goodner, K., Plotto, A., Pritcett, K., and Einstein, M. 2004.
Effect of volatiles and their concentration on perception of tomato
descriptors. Journal of Food Science 69, 310–318.
Baldwin, E.A., Scott, J.W., Shewmaker, C.K., and Schuw, W. 2000. Flavor trivia
and tomato aroma: Biochemistry and possible mechanisms for control
of important aroma components. HortScience 35, 1013–1022.
Bapat, V.A., Trivedi, P.K., Ghosh, A., Sane, V.A., Ganapathi, T.R., and Nath, P.
2010. Ripening of fleshy fruit: Molecular insight and the role of ethyl-
ene. Biotechnology Advances 28, 94–107.
Ben-Arie, R., Segal, N., and Guelfat-Reich, S. 1984. The maturation and ripen-
ing of the ‘Wonderful’ pomegranate. Journal of the American Society for
Horticultural Science 109, 898–902.

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Biale, J.B. 1960. Respiration of fruit. In Ruhland, W. (ed.), Encyclopedia of

Plant Physiology, vol. 12. Springer, Berlin, pp. 536–592.
Biale, J.B. 1976. Recent advances in postharvest physiology of tropical and
subtropical fruits. Acta Horticulturae 57, 179–187.
Biale, J.B., and Barcus, D.E. 1970. Respiratory patterns of tropical fruits of
the Amazon basin. Tropical Science 12, 93–104.
Biale, J.B., and Young, R.E. 1981. Respiration and ripening in fruits—
retrospect and prospect. In Friend, J., and Rhodes, M.J.C. (eds.), Recent
Advances in the Biochemistry of Fruit and Vegetables, 1st ed. Academic
Press, London, pp. 1–39.
Blanpied, G.D. 1972. A study of ethylene in apple, red raspberry and cherry.
Plant Physiology 48, 627–630.
Bouzayen, M., Latche, A., Nath, P., and Pech, J.C. 2010. Mechanism of fruit
ripening. In Pua, E.C., and Davey, M.R. (eds.), Plant Developmental
Biology—Biotechnological Perspectives, vol. 1. Springer-Verlag, Berlin,
pp. 319–339.
Brady, C.J. 1987. Fruit ripening. Annual Review of Plant Physiology and Plant
Molecular Biology 38, 155–178.
Brecht, J., and Yahia, E.M. 2009. Postharvest physiology. In Litz, R. (ed.),
The Mango: Botany, Production and Uses, 2nd ed. CAB International,
Wallingford, UK, pp. 484–528.
Bron, I.U., and Jacomino, A.P. 2006. Ripening and quality of ‘Golden’ papaya
fruit harvested at different maturity stages. Brazilian Journal of Plant
Physiology 18, 389–396.
Brown, A.C., and Summers, W.L. 1985. Carbohydrates accumulation and
color development in watermelon. Journal of the American Society for
Horticulture Science 110, 683–686.
Brown, B.I., and Wills, R.B.H. 1983. Postharvest changes in guava fruits of
different maturity. Scientia Horticulturae 19, 237–243.
Brown, B.I., Wong, I.S., George, A.P., and Nissen, R.J. 1988. Comparative
studies on the postharvest physiology of fruit from different spe-
cies of Annona (custard apple). Journal of Horticultural Science 63,
Brownleader, M.D., Jackson, P., Mobasheri, A., Pantelides, A.T., Sumar, S.,
Trevan, M., and Dey, P.M. 1999. Molecular aspects of cell wall modi-
fications during fruit ripening. Critical Reviews in Food Science and
Nutrition 39, 149–164.
Brummell, D.A. 2006. Cell wall disassembly in ripening fruit. Functional
Plant Biology 33, 103–119.
Brummell, D.A., and Harpster, M.H. 2001. Cell wall metabolism in fruit
softening and quality and its manipulation in transgenic plants. Plant
Molecular Biology 47, 311–340.
Bugaud, C., Alter, P., Daribo, M.O., and Brillouet, J.M. 2009. Comparison of
the physico-chemical characteristics of a new triploid banana hybrid,
FLHORBAN 920, and the Cavendish variety. Journal of the Science of
Food and Agriculture 89, 407–413.


Bugaud, C., Deverge, E., Daribo, M.O., Ribeyre, F., Fils-Lycaon, B., and
Mbéguié-A-Mbéguié, D. 2011. Sensory characterisation enabled the
first classification of dessert bananas. Journal of the Science of Food and
Agriculture 91, 992–1000.
Burdon, J., Dori, S., Marinansky, R., and Pesis, E. 1996. Acetaldehyde inhi-
bition of ethylene biosynthesis in mango fruit. Postharvest Biology and
Technology 8, 153–161.
Burdon, J.N., and Sexton, R. 1990. Fruit abscission and ethylene production
of red raspberry cultivars. Scientia Horticulturae 43, 95–102.
Burg, S.P., and Burg, E.A. 1962. Role of ethylene in fruit ripening. Plant
Physiology 37, 179–189.
Cano, M.P., Ancos, B., Matallana, M.C., Cámara, M., Reglero, G., and
Tabera, J. 1997. Differences among Spanish and Latin American banana
cultivars: Morphological, chemical and sensory characteristics. Food
Chemistry 59, 411–419.
Carrilo-Lopez, A., Cruz-Hernandez, A., Carabez-Trejo, A., Guevara-Lara, F.,
and Paredes-Lopez, O. 2002. Hydrolytic activity and ultrastructural
changes in fruit skins from two prickly pear (Opuntia species) variet-
ies during storage. Journal of Agricultural and Food Chemistry 50,
Carrilo-Lopez, A., Cruz-Hernandez, A., Guevara-Lara, F., and Paredes-Lopez,
O. 2003. Physico-chemical changes during ripening in storage of two
varieties of prickly pear stored at 18°C. Journal of Food Science and
Technology 40, 461–464.
Carrington, C.M.S. 2011. A frim focus on tropical fruit ripening. Acta
Horticulturae 894, 17–32.
Carrington, C.M.S., and King, R.A.G. 2002. Fruit development and ripening
in Barbados cherry (Malpighia emarginata DC.). Scientia Horticulturae
92, 1–7.
Causse, M., Saliba-Colombani, V., Lesschaeve, I., Duffe, P., Rousselle, P., and
Buret, M. 2001. Genetic analysis of organoleptic quality in fresh mar-
ket tomato. 2. Mapping QTLs for sensory attributes. Theoretical and
Applied Genetics 102, 273–283.
Chen, F.X., Liu, X.H., and Chen, L.S. 2009. Developmental changes in
pulp organic acid concentration and activities of acid-metabolising
enzymes during the fruit development of two loquat (Eriobotrya
japonica Lindl.) cultivars differing in fruit acidity. Food Chemistry 114,
Chervin, C., El-Kereamy, A., Roustan, J.P., Latche, A., Lamon, J., and
Bouzayen, M. 2004. Ethylene seems required for the berry develop-
ment and ripening in grape, a non-climacteric fruit. Plant Science 167,
Chervin, C., Tira-umphon, A., Terrier, N., Zouine, M., Severac, D., and
Roustan, J.P. 2008. Simulation of the grape berry expansion by ethyl-
ene and effects on related gene transcripts over the ripening phase.
Physiologia Plantarum 134, 534–546.

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Chyau, C.C., Ko, P.T., Chan, C.H., and Mau, J.L. 2003. Free and glycosidi-
cally bound aroma compounds in lychee (Litchi chinesis Sonn.). Food
Chemistry 80, 387–392.
Clendennen, S.K., and May, G.D. 1997. Differential gene expression in ripen-
ing banana fruit. Plant Physiology 115, 463–469.
Cruz-Hernandez, A., and Paredes-Lopez, O. 2012. Fruit quality: New insights
for biotechnology. Critical Reviews in Food Science and Nutrition 52,
Cua, A.U., and Lizada, M.C.C. 1990. Ethylene production in the ‘Carabao’
mango (Mangifera indica L.) fruit during maturation and ripening. Acta
Horticulturae 269, 169–179.
De Souza, M., Silva-Ferreira, K., Paes-Chaves, J.B., and Lopes-Texeira, S.
2008. L-Ascorbic acid, β-carotein and lycopene in papaya fruits (Carica
papaya) with or without physiological skin frackles. Scientia Agricola
65, 246–250.
de Vasconcelos Facundo, H.V., dos Santos Garruti, D., dos Santos Dias, C.T.,
Cordenunsi, B.R., and Lajolo, F.M. 2012. Influence of different banana
cultivars on volatile compounds during ripening in cold storage. Food
Research International 49, 626–633.
Ding, C.K., Chachin, K., Hamauzu, Y., Ueda, Y., and Imahori, Y. 1998. Effects
of storage temperatures on physiology and quality of loquat fruit.
Postharvest Biology and Technology 14, 309–315.
Ding, P., and Tee, Y.K. 2011. Physicochemical characteristics of dabai
(Canarium odontophyllum Miq.) fruit. Fruits 66, 1–6.
Dixon, J., and Hewett, E.W. 2000. Factors affecting apple aroma/flavour
volatile concentration: A review. New Zealand Journal of Crop and
Horticultural Science 28, 155–173.
Downs, C.G., Brady, C.J., Campbell, J.M., and McGlasson, W.B. 1991. Normal
ripening cultivars of Pyrus serotina are either climacteric or non-climac-
teric. Scientia Horticulturae 48, 213–221.
Drury, R., Hortensteiner, S., Donnison, I., Bird, C.R., and Seymour, G.B.
1999. Chlorophyll catabolism and gene expression in the peel of ripen-
ing banana fruits. Physiologia Plantarum 107, 32–38.
Dumville, J.C., and Fry, S.C. 2003. Solubilisation of tomato fruit pectins by
ascorbate: A possible non-enzymatic mechanism of fruit softening.
Planta 217, 951–961.
Durate, O. 2011. Achachairu (Garcinia humilis (Vahl) C.D. Adam). In
Yahia, E.M. (ed.), Postharvest Biology and Technology of Tropical and
Subtropical Fruits. Woodhead Publishing, Cambridge, UK, pp. 48–53.
Egea, I., Barsan, C., Bian, W., Purgatto, E., Latche, A., Chervin, C., Bouzayen,
M., and Pech, J.C. 2010. Chromoplast differentiation: Current status
and perspectives. Plant and Cell Physiology 51, 1601–1611.
Elkashif, M.E., Huber, D.J., and Brecht, J.K. 1989. Respiration and ethylene
production in harvested watermelon fruit: Evidence for non-climacteric
respiratory behaviour. Journal of the American Society for Horticultural
Science 114, 81–85.


Espley, R.V., Brendolise, C., Chagne, D., Kutty-Amma, S., Green, S., Volz, R.,
Putterill, J., Schouten, H.J., Gardiner, S.E., Hellens, R.P., and Allen, A.C.
2009. Multiple repeats of promoter segments causes transcription factor
autoregulation in red apples. Plant Cell 21, 168–183.
Etienne, A., Génard, M., Lobit, P., Mbeguié-A-Mbéguié, D., and Bugaud, C.
2013. What controls fleshy fruit acidity? A review of malate and citrate
accumulation in fruit cells. Journal of Experimental Botany, doi:10.1093/
Etienne, C., Moing, A., Dirlewanger, E., Raymond, P., Monet, R., and Rothan,
C. 2002. Isolation and characterization of six peach cDNAs encoding
key proteins in organic acid metabolism and solute accumulation:
Involvement in regulating peach fruit acidity. Physiologia Plantarum
114, 259–270.
Fait, A., Hanhineva, K., Beleggia, R., Dai, N., Rogachev, I., Nikiforova, V.J.,
Fernie, A.R., and Aharoni, A. 2008. Reconfiguration of the achene and
receptacle metabolic networks during strawberry fruit development.
Plant Physiology 148, 730–750.
Fanciullino, A.L., Cercos, M., Dhuikue-Mayer, C., Froelicher, Y., Talon, M.,
Ollitrault, P., and Morillon, R. 2008. Changes in carotenoids content
and biosynthetic gene expression in juice sacs of four orange varieties
(Citrus sinensis) differing in flesh fruit color. Journal of Agricultural and
Food Chemistry 56, 3628–3638.
Fidler, J.C., and North, C.J. 1967. The effect of storage on the respiration of
apples. I. The effect of temperature and concentration of CO2 and O2 on
the production of CO2 and uptake of O2. Journal of Horticulture Science
42, 189–206.
Fraser, P.D., Kiano, J.W., Truesdale, M.R., Schuch, W., and Bramley, P.M.
1999. Phytoene synthase-2 enzyme activity in tomato does not contrib-
ute to carotenoid synthesis in ripening fruit. Plant Molecular Biology 40,
Garcia, E., and Lajola, A.M. 1988. Starch transformation during banana ripen-
ing: The amylase and glucosidase behaviour. Journal of Food Science 53,
Gardea, A.A., Martínez-Tellez, M.A., Sanchez, A., Baez, M., and Siller,
J.H. 1994. Postharvest weight loss of Flame Seedless clusters. In
International Symposium on Table Grape Production, Anaheim, CA,
pp. 203–206.
Gariglio, N., Juan, M., Castillo, A., Almela, V., and Agustí, M. 2002.
Morphological, histological and physiological study of purple spot of
loquat fruit. Scientia Horticulturae 92, 255–263.
Gariglio, N.F., Reig, C., and Agusti, M. 2008. Assimilate partitioning between
the flesh and the rind is responsible for purple spot in loquat fruit.
Journal of Horticultural Science and Biotechnology 83, 37–42.
Giovannoni, J. 2001. Molecular biology of fruit maturation and ripening.
Annual Review of Plant Physiology and Plant Molecular Biology 52,

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Gonzalez, L., Lafuente, M.T., and Zacarias, L. 2004. Maturation of loquat fruit
(Eriobotrya japonica Lindl.) under Spanish growing conditions and its
postharvest performance. Options Mediterran 58, 171–179.
Goulao, L.F., and Oliveira, C.M. 2008. Cell wall modifications during fruit rip-
ening: When a fruit is not fruit. Trends in Food Science and Technology
19, 4–25.
Graham, O.S., Mohammed, M., and Wickham, L.D. 2004. Effects of heat
treatments on the quality of miniature golden apples (Spondias cytherea
Sonn) during low temperature storage. International Journal of Food,
Agriculture and Environment 2, 43–48.
Gurrieri, F., Audergon, J.M., Albagnac, G., and Reich, M. 2001. Soluble sug-
ars and carboxylic acids in ripe apricot fruit as parameters for distin-
guishing different cultivars. Euphytica 117, 183–189.
Hamauzu, Y., Chachin, K., Ding, C.K., and Kurooka, H. 1999. Differences in
surface color, flesh firmness, physiological activity, and some compo-
nents of loquat fruit picked at various stages of maturity. Journal of the
Japanese Society for Horticultural Science 65, 859–865.
Hardy, P.J. 1968. Metabolism of sugars and organic acids in immature grape
berries. Plant Physiology 43, 224–228.
Hasegawa, P.N., Faria, A.F., Mercadante, A.Z., Chagas, E.A., Pio, R., Lajolo,
F.M., Cordenunsi, B.R., and Purgatto, E. 2010. Chemical composition of
five loquat cultivars planted in Brazil. Ciencia e Tecnologia de Alimentos
30, 552–559.
Hernandez, M.S., Barrera, J.A., Martinez, O., and Fernandez-Trujillo, J.P.
2009. Postharvest quality of araza fruit during low temperature storage.
LWT Food Science and Technology 42, 879–884.
Hernandez, M.S., Martinez, O., and Fernandez-Trujillo, J.P. 2007. Behavior
of araza fruit quality traits during growth, development and ripening.
Scientia Horticulturae 111, 220–227.
Hirai, M. 1980. Sugar accumulation and development of loquat fruit. Journal
of the Japanese Society of Horticultural Science 49, 347–353.
Huber, D.J. 1984. Strawberry fruit softening: The potential roles of polyur-
onoids and hemicelluloses. Journal of Food Science 49, 1310–1315.
Hulme, A.C. 1958. Some aspects of the biochemistry of apple and pear fruit.
Advances in Food Research 8, 297–413.
Hulme, A.C. 1970. The Biochemistry of Fruits and Their Products, vol. I.
Academic Press, London, p. 620.
Ilahy, R., Hdider, C., Lenucci, M.S., Tlili, I., and Dalessandro, G. 2011.
Antioxidant activity and bioactive compound changes during fruit ripen-
ing of high-lycopene tomato cultivars. Journal of Food Composition and
Analysis 24, 588–595.
Ishimaru, M., and Kobayashi, S. 2002. Expression of a xyloglucan endo-
transglycosilase gene is closely related to grape berry softening. Plant
Science 162, 621–628.
Ismael, A.A., and Render, W.T. 1969. Evidence of a respiratory climacteric in
highbush and lowbush blueberry fruit. HortScience 4, 342–344.


Itai, A., Tanabe, K., Tamura, F., and Tanaka, T. 2000. Isolation of cDNA clones
corresponding to genes expressed during fruit ripening in Japanese
pear (Pyrus pyrifolia Nakai): involvement of the ethylene signal trans-
duction pathway in their expression. Journal of Experimental Botany 51,
Iwaoka, W., Hagi, Y., Umano, K., and Shibamoto, T. 1994. Volatile chemicals
identified in fresh and cooked breadfruit. Journal of Agricultural and
Food Chemistry 42, 975–976.
Jiang, J.P., Su, M.X., and Lee, P.M. 1986. The production and physiological
effects of ethylene during ontogeny and after harvest of litchi fruits.
Acta Phytophysiologia Sinica 12, 95–103.
Jiang, W.B., and Sheng, Q. 2003. Effects of 1-methylcyclopropene and giberel-
lic acid on ripening of jujube (Zizyphus jujuba M) in relation to quality.
Journal of the Science of Food and Agriculture 84, 31–35.
Jordan, M.J., Tandon, K., Shaw, P.E., and Goodner, K.L. 2001. Aromatic profile
of aqueous banana essence and banana fruit by gas ­chromatography–
mass spectrometry (GC–MS) and gas chromatography–olfactometry
(GC–O). Journal of Agricultural and Food Chemistry 49, 4813–4817.
Joyce, D.C., and Li, J.R. 2003. Postharvest characteristics of red bayberry.
In Proceedings of the Australasian Postharvest Horticulture Conference,
Brisbane, Australia, October 1–3, pp. 125–226.
Kader, A. 2002. Loquat: recommendations for maintaining postharvest qual-
Kader, A.A., and Arpaia, M.L. 2001. Avocado: recommendations for main-
taining postharvest quality.­
producefacts/fruit/ avocado.html.
Kader, A.A., Li, Y., and Chordas, A. 1982. Postharvest respiration, ethyl-
ene production, and compositional changes in Chinese jujube fruits.
HortScience 17, 678–679.
Kalra, S.K., and Tandon, D.K. 1983. Ripening behavior of Dashehari mango in
relation to harvest period. Scientia Horticulturae 19, 263–269.
Katz, E., Lagunes, P.M., Riov, J., Weiss, D., and Goldschmidt, E.E. 2004.
Molecular and physiological evidence suggests the existence of a sys-
tem II-like pathway of ethylene production in non-climacteric citrus
fruit. Planta 219, 243–252.
Kays, S. 1991. Postharvest Physiology of Perishable Plant Products. Van
Nostrand Reinhold, New York.
Kays, S.J., and Hayes, M.J. 1978. Induction of ripening in the fruits of bitter
gourd by ethylene. Tropical Agriculture 55, 167–171.
Ketsa, S., and Daengkanit, T. 1998. Physiological changes during posthar-
vest ripening of durian fruit (Durio zibethinus Murray). Journal of
Horticultural Science and Biotechnology 73, 575–577.
Klee, H.J. 2002. Control of ethylene-mediated processes in tomato at the level
of receptors. Journal of Experimental Botany 53, 2057–2063.

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Kliewer, W.M. 1971. The effect of day temperature and light intensity on con-
centration of malic and tartaric acids in Vitis vinifera L. grapes. Journal
of the American Society for Horticultural Science 96, 372–377.
Kobayashi, S., Goto-Yamamoto, N., and Hirochika, H. 2004. Retrotransposon-
induced mutations in grape skin color. Science 304, 982.
Krishnamurthy, S., and Subramanyam, H. 1970. Respiratory climacteric and
chemical changes in the mango fruit Mangifera indica L. Journal of the
American Society for Horticultural Science 95, 333–337.
Kurz, C., Carle, R., and Schieber, A. 2008. HPLC-DAD-MS characterization
of carotenoids from apricots and pumpkins for the evaluation of fruit
product authenticity. Food Chemistry 110, 522–530.
Lakshminarayana, S., and Estrella, I.B. 1978. Postharvest respiratory behav-
iour of tuna (prickly pear) fruit (Opuntia robusta Mill.). Journal of
Horticultural Science 53, 327–330.
Lam, P.F., and Wan, C.K. 1983. Climacteric nature of the carambola (Averrhoa
carambola L.) fruit. Pertanika 6, 44–47.
Lannetta, P.P.M., Laarhoven, L.I., Medina-Escobar, N., James, E.K., McManus,
M.T., Davies, H.V., and Harren, F.J.M. 2006. Ethylene and carbon diox-
ide production by developing strawberries show a correlative pattern
that is indicative of ripening climacteric fruit. Physiologia Plantarum
127, 247–259.
Lazan, H., Ali, Z.M., Soh, J.S., and Talkah, Z. 1993. The biochemical basis of
differential ripening in mango. Acta Horticulturae 341, 500–509.
Lelievre, J.M., Latche, A., Jones, B., Bouzayen, M., and Pech, J.C. 1997.
Ethylene and fruit ripening. Physiologia Plantarum 101, 727–739.
Li, X., Xu, C., Korban, S.S., and Chen, K. 2010. Regulatory mechanisms of
textural changes in ripening fruits. Critical Reviews in Plant Sciences
29, 222–243.
Lipe, J.A. 1978. Ethylene in fruits of blackberry and rabbiteye blueberry.
Journal of the American Society for Horticultural Science 103, 76–77.
Lo Bianco, R., and Rieger, M. 2002. Partitioning of sorbitol and sucrose catab-
olism within peach fruit. Journal of the American Society for Horticulture
Science 127, 115–121.
Lu, X.-P., Liu, Y.-Z., Zhou, G.-F., Wei, Q.-J., Hu, H.-J., and Peng, S.-A. 2011.
Identification of organic acid-related genes and their expression profiles
in two pear (Pyrus pyrofolia) cultivars with difference in predominant
acid type at fruit ripening stage. Scientia Horticulturae 129, 680–687.
Lyons, J.M., McGlasson, W.B., and Pratt, H.K. 1962. Ethylene production,
respiration and internal gas concentrations in cantaloupe fruits at vari-
ous stages of maturity. Plant Physiology 37, 31–36.
Malik, S.K., Kumar, S., and Bansal, K.C. 2014. Genetic diversity of tropical
fruit. In Nath, P., Bouzayen, M., Mattoo, A.K., and Pech, J.C. (eds.),
Fruit Ripening: Physiology, Signaling and Genomics. CABI, Oxfordshire,
UK, pp. 217–227.


Manenoi, A., Bayogan, E.R.V., Thumdee, S., and Paull, R.E. 2007. Utility of
1-methylcyclopropene as a papaya postharvest treatment. Postharvest
Biology and Technology 44, 55–62.
Marei, N., and Crane, J.C. 1971. Growth and respiratory response of fig
(Ficus  carica L. cv. Mission) fruits to ethylene. Plant Physiology 48,
Marin-Rodriguez, M.C., Orchard, J., and Seymour, G.B. 2002. Pectate lyases,
cell wall degradation and fruit softening. Journal of Experimental Botany
53, 2115–2119.
Mattoo, A.K., and Modi, V.V. 1969. Biochemical aspects of ripening and chill-
ing injury in mango fruit. In Proceedings of International Conference on
Tropical and Subtropical Fruit, London, pp. 111–115.
Maxie, E.C., Catlin, P.B., and Hartman, H.T. 1960. Respiration and ripening of
olive fruits. Proceedings of the American Society for Horticultural Science
75, 275–291.
McCarthy, A.I., Palmer, J.K., Shaw, C.P., and Anderson, E.E. 1963. Correlation
of gas chromatographic data for flavour profiles of fresh banana fruits.
Journal of Food Science 28, 379–384.
Medina-Suarez, R., Manning, K., Fletcher, J., Aked, J., Bird, C.R., and
Seymour, G.B. 1997. Gene expression in the pulp of ripening bananas
(two-dimensional sodium dodecyl sulfate-polyacrylamide gel electro-
phoresis of in vitro translation products and cDNA cloning of 25 differ-
ent ripening-related mRNAs). Plant Physiology 115, 453–461.
Mendoza, D.B., Pantastico, E., and Javier, F.B. 1972. Storage and handling
of rambutan (Nephelium lappaceum L.). Philippines Agriculture 55,
Moore, S., Vrebalov, J., Payton, P., and Giovannoni, J. 2002. Use of genom-
ics tools to isolate key ripening genes and analyse fruit maturation in
tomato. Journal of Experimental Botany 53, 2023–2030.
Morandi, B., Grappadelli, L.G., Rieger, M., and Lo Bianco, R. 2008.
Carbohydrate availability affects growth and metabolism in peach fruit.
Physiologia Plantarum 133, 229–241.
Mota, W.F., Salomao, L.C.C., Perira, M.C.T., and Cecon, P.R. 2002. The influ-
ence of the postharvest treatment with calcium in jaboticaba fruits con-
servation. Revista Brasieira de Fruticultura 24, 49–52.
Nam, Y.W., Tichit, L., Leperlier, M., Cuerq, B., Marty, I., and Lelievre, J.M.
1999. Isolation and characterization of mRNA differentially expressed
during ripening of wild strawberry (Fragaria vesca L.) fruits. Plant
Molecular Biology 39, 629–636.
Nerd, A., Gutman, F., and Mizrahi, Y. 1999. Ripening and postharvest behav-
iour of fruits of two Hylocereus species (Cactaceae). Postharvest Biology
and Technology 17, 39–45.
Nerd, A., and Mizrahi, Y. 1999. The effect of ripening stage on fruit quality
after storage of yellow pitaya. Postharvest Biology and Technology 15,

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Nogueira, J.M., Fernandes, P.J., and Nascimento, A.M. 2003. Composition

of volatiles of banana cultivars from Madeira Island. Phytochemical
Analysis 14, 87–90.
Nunan, K.J., Davies, C., Robinson, S.P., and Fincher, G.B. 2001. Expression
patterns of cell wall modifying enzymes during grape berry develop-
ment. Planta 214, 257–264.
Oslund, C., and Davenport, T. 1983. Ethylene and carbon dioxide in ripening
fruit of Averrhoa carambola. HortScience 18, 229–230.
Palma, J.M., Corpas, F.J., and del Rio, L.A. 2011. Proteomics as an approach
to the understanding of the molecular physiology of fruit development
and ripening. Journal of Proteomics 74, 1230–1243.
Pareek, S., and Kitinoja, L. 2011. Aonla (Emblica officinalis Gaertn.). In
Yahia, E.M. (ed.), Postharvest Biology and Technology of Tropical
and Subtropical  Fruits, Woodhead Publishing Ltd., Cambridge, UK,
pp. 65–97.
Pareek, S., and Yahia, E.M. 2013. Postharvest biology and technology of ber
fruit. Horticultural Reviews 41, 201–240.
Pareek, S., Benkeblia, N., Janick, J., Cao, S., and Yahia, E.M. 2014. Postharvest
physiology and technology of loquat (Eriobotria japonica Lindl.) fruit.
Journal of the Science of Food and Agriculture 94, 1495–1504.
Pareek, S., Kitinoja, L., Kaushik, R.A., and Paliwal, R. 2009. Postharvest physi-
ology and storage of ber. Stewart Postharvest Review 5, 1–10.
Paul, V., Pandey, R., and Srivastava, G.C. 2012. The fading distinctions
between classical patterns of ripening in climacteric and non-climacteric
fruit and the ubiquity of ethylene: an overview. Journal of Food Science
and Technology 49, 1–21.
Paull, R.E., and Ketsa, S. 2004. Mangosteen. In Gross, K.C., Wang, C.Y.,
and Saltveit, M.E. (eds.), The Commercial Storage of Fruits, Vegetables
and Florists and Nursery Stocks. USDA Handbook No. 66 revised. U.S.
Department of Agriculture, Agricultural Research Service, Washington,
DC. (accessed
February 23, 2015).
Payasi, A., and Sanwal, G.G. 2005. Biochemistry of fruit ripening. Indian
Journal of Agricultural Biochemistry 18, 51–60.
Payasi, A., and Sanwal, G.G. 2009. Molecular mechanism of ethylene signal
transduction in fruits. Indian Journal of Agricultural Biochemistry 22,
Pech, J.C., Bouzayen, M., and Latche, A. 2008. Climacteric fruit ripening:
ethylene-dependent and independent regulation of ripening pathways
in melon fruit. Plant Science 175, 114–120.
Pech, J.C., Purgatto, E., Bouzayen, M., and Latche, A. 2012. Ethylene and
fruit ripening. Annual Plant Reviews 44, 275–304.
Pech, J.C., Purgatto, E., Girardi, C.L., Rombaldi, C.V., and Latche, A. 2013.
Current challenges in postharvest biology of fruit ripening. Current
Agricultural Science and Technology 19, 1–18.


Pérez, A.G., Cert, A., Rios, J.J., and Olias, J.M. 1997. Free and glycosidi-
cally bound volatile compounds from two banana cultivars: Valery
and Pequena Enana. Journal of Agricultural and Food Chemistry 45,
Perkins-Veazie, P., and Nonneeke, G. 1992. Physiological changes during rip-
ening of raspberry fruit. HortScience 27, 331–333.
Perkins-Veazie, P., Collins, J.K., Pair, S.D., and Roberts, W. 2001. Lycopene
content differs among red-fleshed watermelon cultivars. Journal of the
Science of Food and Agriculture 81, 983–987.
Pontes, M., Pereira, J., and Câmara, J.S. 2012. Dynamic headspace solid-
phase microextraction combined with one-dimensional gas chroma-
tography-mass spectrometry as a powerful tool to differentiate banana
cultivars based on their volatile metabolite profile. Food Chemistry 134,
Pratt, H.K., and Goeschl, J.D. 1968. The role of ethylene in fruit ripening. In
Wightman, F., and Setterfield, G. (eds.), Biochemistry and Physiology of
Plant Growth Substances. Runge, Ottawa, pp. 1295–1302.
Pratt, H.K., and Mendoza, D.B. 1980. Fruit development and ripening of the
star apple (Chrysophyllum cainito L.). HortScience 15, 721–722.
Pratt, H.K., and Reid, M.S. 1974. Chinese gooseberry: Seasonal patterns in
fruit growth and maturation, ripening, respiration and role of ethylene.
Journal of the Science of Food and Agriculture 25, 747–753.
Pratt, H.K., and Reid, M.S. 1976. The tamarillo: Fruit growth and maturation,
ripening, respiration, and the role of ethylene. Journal of the Science of
Food and Agriculture 27, 399–404.
Prinsi, B., Negri, A.S., Fedeli, C., Morgutti, S., Negrini, N., Cocucci, M., and
Espen, L. 2011. Peach fruit ripening: A proteomic comparative analysis
of the mesocarp of two cultivars with different flesh firmness at two
ripening stages. Phytochemistry 72, 1251–1262.
Reid, M.S. 1975. The role of ethylene in the ripening of some unusual fruits.
In Facteurs et regulation de la maturation des fruits, no. 238. CNRS,
Paris, pp. 177–182.
Reig, C., Martinez-Fuentes, A., Juan, M., Gariglio, N., Marti, G., Mesejo, C.,
and Agusti, M. 2007. Tecnicasparaanticipar la recoleccion del fruto del
nisperojapones (Eriobotrya japonica Lindl.). In XI Congress National
SECH, Spain, abstract 4D01.
Rogiers, S.Y., Mohan-Kumar, G.N., and Knowles, N.R. 1998. Regulation of
ethylene production and ripening by Saskatoon (Amelanchier alnifolia
Nutt.) fruit. Canadian Journal of Botany 76, 1743–1754.
Roy, S.K. 1975. Studies of bael fruit (Aegle marmelos Carrea)—changes in
constituents during development, ripening and storage and process-
ing of the fruit. PhD thesis, Bidhan Chandra Krishi Vishva Vidyalaya,
Kalyani, West Bengal, India.
Rygg, G.L. 1975. Date Development, Handling and Packing in the United
States. Agricultural Handbook No. 482. U.S. Department of Agriculture,
Agricultural Research Service, Washington, DC, p. 56.

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Salmon, B., Martin, G.J., Remaud, G., and Fourel, F. 1996. Compositional and
isotopic studies of fruit flavours. Part I. The banana aroma. Flavour and
Fragrance Journal 11, 353–359.
Saltveit, M.E. 1977. Carbon dioxide, ethylene, and color development in rip-
ening mature green bell peppers. Journal of the American Society for
Horticultural Science 102, 523–525.
Sampaio, S.A., Bora, P.S., Holschuh, H.J., and Silva, S.M. 2007. Postharvest
respiratory activity and changes in some chemical constituents dur-
ing maturation of yellow mombin (Spondias mombin) fruit. Ciencia e
Technologia de Alimentos 27, 1–9.
Santos, A.F. dos, Silva, S.M., and Alves, R.E. 2006. Armezenamento de pitanga
sob atmosfera modificada refrigeracao: I-transformacoes quimicas em
pos-colheita. Revista Brasieira de Fruticultura 28, 36–41.
Saradhuldhat, P., and Paull, R.E. 2007. Pineapple organic acid metabolism
and accumulation during fruit development. Scientia Horticulturae 112,
Selvaraj, Y., and Pal, D.K. 1989. Biochemical changes during the ripening
of jackfruit (Artocarpus heterophyllus L.). Journal of Food Science and
Technology 25, 304–307.
Seymour, G.B., Manning, K., Eriksson, E.M., Popovich, A.H., and King, G.J.
2002. Genetic identification and genomic organization of factors affect-
ing fruit texture. Journal of Experimental Botany 5, 2065–2071.
Shen, L., Sheng, J.P., Niu, J.S., and Liu, Q.X. 2004. Changes of respiration
and ethylene production and effects of 1-MCP during the fermentation
softening of Chinese winter jujube fruit. Journal of China Agricultural
University 9, 36–39.
Shiota, H. 1993. New esteric compounds in the volatiles of banana fruit (Musa
sapientum L.). Journal of Agricultural and Food Chemistry 41, 2056–2062.
Sinesio, F., Cammareri, M., Moneta, E., Navez, B., Peparaio, M., Causse,
M., and Grandillo, S. 2010. Sensory quality of fresh French and Dutch
market tomatoes: A preference mapping study with Italian consumers.
Journal of Food Science 75, 55–67.
Singh, S.P., and Pal, R.K. 2008. Response of climacteric-type guava (Psidium
guajava L.) to postharvest treatment with 1-MCP. Postharvest Biology
and Technology 47, 307–314.
Siriphanich, J., Abdulla, H., Kosittrakun, M., and Lizada, M.C.C. 1994.
Physiology. In Nanthachai, S. (ed.), Durian Fruit Development,
Postharvest Physiology, Handling and Marketing in ASEAN. ASEAN
Food Handling Bureau, Kuala Lumpur, pp. 48–57.
Sivakumar, D., Naudé, Y., Rohwer, E., and Korsten, L. 2008. Volatile com-
pounds, quality attributes, mineral composition and pericarp structure
of South African litchi export cvs. Mauritius and McLean’s Red. Journal
of the Science of Food and Agriculture 88, 1074–1081.
Sweetman, C., Deluc, L.G., Cramer, G.R., Ford, C.M., and Soole, K.L. 2009.
Regulation of malate metabolism in grape berry and other developing
fruits. Phytochemistry 70, 1329–1344.


Thompson, A.J., Tor, M., Barry, C.S., Vrebalov, J., Orfila, C., Jarvis, M.C.,
Giovannoni, J.J., Grierson, D., and Seymour, G.B. 1999. Molecular and
genetic characterization of a novel pleiotropic tomato-ripening mutant.
Plant Physiology 120, 383–389.
Tian, M., Sheng, C.C., and Li, Y. 1987. Changes of ethylene synthesis, activity
of polyphenol oxidase, permeability of plasma membrane of Dutch pear
at low temperature. Acta Botanic Sinica 29, 614–619.
Toivonen, P.M.A., and Brummell, D.A. 2008. Biochemical bases of appear-
ance and texture changes in fresh-cut fruit and vegetables. Postharvest
Biology and Technology 48, 1–14.
Tongdee, S.C., Suwanakul, A., and Neamprem, S. 1990. Durian fruit ripen-
ing and the effect of variety, maturity stage at harvest, and atmospheric
gases. Acta Horticulturae 269, 323–334.
Topuz, A., and Ozdemir, F. 2007. Assessment of carotenoids, capsaicinoids
and ascorbic acid composition of some selected pepper cultivars
(Capsicum annuum L.) grown in Turkey. Journal of Food Composition
and Analysis 20, 596–602.
Tressl, R., and Jennings, W.G. 1972. Production of volatile compounds in
the ripening banana. Journal of Agricultural and Food Chemistry 20,
Trinchero, G.D., Sozzi, G.O., Cerri, A.M., Vilella, F., and Fraschina, A.A. 1999.
Ripening-related changes in ethylene production, respiration rate and
cell-wall enzyme activity in goldenberry (Physalis peruviana L.), a sola-
naceous species. Postharvest Biology and Technology 16, 139–145.
Tucker, G.A. 1993. Introduction. In Seymour, G.B., Taylor, J.E., and Tucker,
G.A., Biochemistry of Fruit Ripening. Springer, London, UK. pp. 1–52.
Tucker, G.A., and Grierson, D. 1987. Fruit ripening. In Davies, D. (ed.), The
Biochemistry of Plants, vol. 12. Academic Press, New York, pp. 265–319.
Ulrich, D., Hoberg, E., Rapp, A., and Kecke, S. 1997. Analysis of strawberry
flavor—discrimination of aroma types by quantification of volatile com-
pounds. Zeitschrift fur Lebensmittel-Untersuchung und Forschung A 205,
Velasquez de Klimo, I. 2006. Manejo pos cosecha del nance (Byrsonima crassi-
folia (L.) HBK). Ministerio de Agricultura y Ganaderia, IICA Frutales,
Programa Nacional de Frutales, San Salvador, El Salvador.
Vermeir, S., Hertog, M.L.A.T.M., Vankerschaver, K., Swennen, R., Nicolai,
B.M., and Lammertyn, J. 2009. Instrumental based flavour characteriza-
tion of banana fruit. LWT Food Science and Technology 42, 1647–1653.
Vicente, A.R., Ortugno, C., Powell, A.L., Greve, L.C., and Labavitch, J.M.
2007a. Temporal sequence of cell wall disassembly events in developing
fruits. 1. Analysis of raspberry (Rubus idaeus). Journal of Agricultural
and Food Chemistry 55, 4119–4124.
Vicente, A.R., Ortugo, C., Rosli, H., Powell, A.L., Greve, L.C., and Labavitch,
J.M. 2007b. Temporal sequence of cell wall disassembly events in devel-
oping fruits. 2. Analysis of blueberry (Vaccinium species). Journal of
Agricultural and Food Chemistry 55, 4125–4130.

Ripen in g P hy s i o l o g y : A n Ov e r v i e w

Vicente, A.R., Powell, A., Greve, L.C., and Labavitch, J.M. 2007c. Cell wall disas-
sembly events in boysenberry (Rubus idaeus L. × Rubus ursinus Cham. &
Schldl.) fruit development. Functional Plant Biology 34, 614–623.
Vicente, A.R., Saladie, M., Rose, J.K.C., and Labavitch, J.M. 2007d. The link-
age between cell wall metabolism and fruit softening: looking to the
future. Journal of the Science of Food and Agriculture 87, 1435–1448.
Vrebalov, J., Ruezinsky, D., Padmanabhan, V., White, R., Medrano, D., Drake, R.,
Schuch, W., and Giovannoni, J. 2002. A MADS-box gene necessary for fruit
ripening at the tomato ripening-inhibitor (rin) locus. Science 296, 343–346.
Wade, N.L., Kavanagh, E.E., Hockley, D.G., and Brady, C.J. 1992. Relationship
between softening and the polyuronides in ripening banana fruits.
Journal of the Science of Food and Agriculture 60, 61–68.
Walsh, C.S., Popenoe, J., and Solomos, T. 1983. Thornless blackberry is a
­climacteric fruit. HortScience 18, 482–483.
Walker, A.R., Lee, E., Bogs, J., McDavid, D.A.J., Thomas, M.R., and Robinson,
S.P. 2007. White grapes arose through the mutation of two similar and
adjacent regulatory genes. Plant Journal 49, 772–785.
Wang, T., Gonzalez, A.R., Gbur, E.E., and Aselage, J.M. 1993. Organic acid
changes during ripening of processing peaches. Journal of Food Science
58, 631–632.
Warren, O. 2009. Quality of carambola fruit (Averrhoa carambola L.) as
affected by harvest maturity, postharvest wax coating, ethylene, and
1-methylcyclopropene. MSc thesis, Horticultural Sciences Department,
University of Florida, Gainesville.
Waters, D.L.E., Holton, T.A., Albett, E.M., Lee, L.S., and Henry, R.J. 2005.
cDNAs microarrays analysis of developing grape (Vitis vinifera cv.
Shiraz) berry skin. Functional and Integrative Genomics 5, 40–58.
Watkins, C.B. 2002. Ethylene biosynthesis: Mode of action, consequences,
and control. In Knee, M. (ed.), Fruit Quality and Its Biological Basis,
Sheffield Academic Press, Sheffield, pp. 180–224.
Watson, R., Wright, C.J., McBurney, T., Taylor, A.J., and Linforth, R.S.T. 2002.
Influence of harvest date and light integral on the development of straw-
berry flavour compounds. Journal of Experimental Botany 53, 2121–2129.
Weaver, R.J., and Singh, I.S. 1978. Occurrence of endogenous ethylene and
effect of plant regulators on ethylene production in the grapevine.
American Journal of Enology and Viticulture 29, 282–285.
White, P.J. 2002. Recent advances in fruit development and ripening: An over-
view. Journal of Experimental Botany 53, 1995–2000.
Whiting, G.C. 1970. Sugars. In Hulme, A.C. (ed.), The Biochemistry of Fruits
and Their Products, vol. I. Academic Press, London, pp. 1–620.
Wu, Y., Pan, Q., Qu, W., and Duan, C. 2009. Comparison of volatile profiles
of nine litchi (Litchi chinensis Sonn.) cultivars from southern China.
Journal of Agricultural and Food Chemistry 57, 9676–9681.
Yahia, E.M. 2004. Sapodilla and related fruits. In Gross, K.C., Wang, C.Y.,
and Saltveit, M.E. (eds.), The Commercial Storage of Fruits, Vegetables
and Florists and Nursery Stocks. USDA Handbook No. 66 revised.


U.S.  Department of Agriculture, Agricultural Research Service,

Washington, DC.
(accessed February 23, 2015).
Yamaguchi, S., Yoshikawa, T., Ikeda, S., and Ninomiya, T. 1970. Studies on
the taste of some sweet substances. Part I. Measurement of the relative
sweetness. Agricultural and Biological Chemistry 34, 181–186.
Yamaki, S. 1984. Isolation of vacuoles from immature apple fruit flesh and
compartmentation of sugars, organic acids, phenolic compounds and
amino acids. Plant and Cell Physiology 25, 151–166.
Yamaki, Y.T. 1989. Organic acids in the juice of citrus fruits. Journal of the
Japanese Society for Horticultural Science 58, 587–594.
Yang, L.T., Xie, C.Y., Jiang, H.X., and Chen, L.S. 2011. Expression of six
malate-related genes in pulp during the fruit development of two loquat
(Eriobotrya japonica) cultivars differing in fruit acidity. African Journal
of Biotechnology 10, 2414–2422.
Yelle, S., Chetelat, R.T., Dorais, M., Deverna, J.W., and Bennett, A.B. 1991.
Sink metabolism in tomato fruit. IV. Genetic and biochemical analysis
of sucrose accumulation. Plant Physiology 95, 1026–1035.
Zhang, X., Yazaki, J., Sundaresan, A., Cokus, S., Chan, S., Chen, H., Henderson,
I., Shinn, P., Pellegrini, M., Jacobsen, S., and Ecker, J.R. 2006. Genome-
wide high resolution mapping and functional analysis of DNA methyla-
tion in Arabidopsis. Cell 126, 1189–1201.
Zhao, Y.F., Lin, H.T., Lin, J.F., Chen, S.J., and Xi, Y.F. 2005. Changes of respi-
ration rate, cell membrane permeability and quality in postharvest lon-
gan fruit. Journal of Fujian Agriculture and Forest University (Natural
Science Edition) 34, 263–268.
Zhou, C.H., Xu, C.J., Sun, C.D., Li, X., and Chen, K.S. 2007. Carotenoids in
white- and red-fleshed loquat fruits. Journal of Agricultural and Food
Chemistry 55, 7822–7830.

Chapter 2

Physiology of Fruits
and Vegetables
Peter M.A. Toivonen
Agriculture and Agri-Food Canada
Summerland, British Columbia, Canada

Abstract 50
2.1 Introduction 51
2.2  Classification of Fruits and Vegetables Based on
Physiological Characteristics 51
2.2.1  Ethylene Biology 51
2.2.2  Respiratory Characteristics: Climacteric and
Nonclimacteric 56
2.2.3  Developmental or Maturity Stage at Harvest 56
2.2.4  Tolerance to Low Temperatures 58
2.3  Factors Affecting Physiology of Fruits and Vegetables in
Postharvest Systems 60
2.3.1 Temperature 60
2.3.2 Humidity 62
2.3.3  Atmospheric Modification 64
2.3.4  Abiotic Stresses 65


2.4  Physiological Changes Occurring during Postharvest

Handling or Storage 66
2.4.1  Depletion of Respiratory Substrate 66
2.4.2  Hormonal Effects 68
2.4.3  Membrane Alterations 71
2.5  Conclusions and Future Perspectives 71
References 72

This chapter covers a wide range of issues pertaining to the many aspects
of fruit and vegetable tissue physiology and changes that occur during
postharvest handling and storage. Understanding the specific ethyl-
ene biology, respiration behavior, and developmental or maturity stage
at harvest can provide great insight to postharvest quality retention
characteristics of a fruit or vegetable. Ethylene itself is one of the most
important factors determining quality retention, with different fruit and
vegetable classes having differing production rates and sensitivity to the
levels of ethylene present. Numerous disorders and quality defects can
be attributed to exposures to ethylene in postharvest systems, and these
are discussed. Respiration behaviors can be separated into two classes:
(1) climacteric and (2) nonclimacteric. The classification of respira-
tory behavior will have an impact on postharvest approaches required
for optimal handling of a particular fruit or vegetable in question. The
anatomical structure and developmental maturity of a fruit or vegetable
can range from an immature vegetable tissue all the way to a ripe fruit
tissue, and this will also determine which postharvest procedures will
be successful for handling and storage life potential. The importance of
temperature, humidity, and atmosphere management on quality reten-
tion is discussed in a mechanistic manner to explain the physiological
responses to these postharvest interventions. Finally, endogenous and
exogenously applied phytohormones are important to signaling meta-
bolic shifts that influence quality retention, and these implications are
explored. The discussion in this chapter is intended to provide a basic
and mechanistic understanding of how and why quality changes occur
in postharvest handling of fruits and vegetables that have contrasting
physiological characteristics. The discussion culminates in discussion
on improving quality retention of fruits and vegetables through multi-
disciplinary cooperation of plant breeders, physiologists, biochemists,
and molecular biologists, leading to more robust solutions to postharvest

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

2.1 Introduction
Development of successful harvest, handling, storage, and distribution
protocols for fruits and vegetables has relied on the understanding of the
physiology of each type of produce and how that physiology responds to
the postharvest environment to which it is exposed (Toivonen and Hodges,
2011). Because of limitations in produce handling systems, many different
types of fruits and vegetables are held in common rooms or containers,
and this has led to issues of incompatibility of produce having different
physiologies. This incompatibility is expressed as shorter storage or shelf
life, development of off-flavors, and accumulation of bitter compounds
(Thompson et al., 1996). Another consequence of common storage areas
is that some fruits and vegetables have differing tolerances to low tempera-
tures, and therefore some fruits or vegetables held in common-use, low-
temperature storage might suffer chilling injuries (Thompson et al., 1996;
Cantwell, 2002). This phenomenon continues to exist and can still today
be visualized in retail produce displays as pitting with associated decay in
cantaloupes and summer squash or as sheet pitting in green peppers (per-
sonal observation). The challenge facing postharvest science and technol-
ogy today is the development of innovative strategies to resolve problems
associated with differing fruit and vegetable physiologies in the real-world
harvest, storage, distribution, and retail systems.
This chapter is devoted to providing an overview of the fundamen-
tal aspects of fruit and vegetable postharvest physiologies to allow a better
appreciation for challenges faced in their handling. Understanding of the
fundamental principles will provide grounded principles that will hopefully
be the basis of new, innovative strategies for harvest, storage, handing, and
distribution of these living and very perishable products.

2.2  Classification of Fruits and Vegetables

Based on Physiological Characteristics
There are a number of aspects of fruit or vegetable physiology that can
influence postharvest quality outcomes. These aspects include ethylene
biology, respiration, the developmental or maturity stage at harvest, and
sensitivity to low temperature.

2.2.1  Ethylene Biology

Ethylene is arguably the most important molecule in the postharvest han-
dling of fruits and vegetables, having a wide range of effects that lead to qual-
ity loss (Saltveit, 1999). The reason for ethylene’s prominence on postharvest


­ andling is that it is a plant growth regulator that is naturally produced with

ripening of climacteric fruit and generally in response to either abiotic or
biotic stresses (Saltveit, 1999). However, another feature of postharvest han-
dling is that fruits and vegetables are usually held in containers, chambers,
or rooms, and this allows any ethylene produced by the produce to accumu-
late. Quite often, fruits and vegetables that produce ethylene or are sensi-
tive to ethylene are handled alongside each other, leading to quality losses
in sensitive produce in that containment (Saltveit, 1999). While thresholds
to ethylene sensitivity have been generally thought to be around 0.1 µl L –1,
there is evidence suggesting that much lower levels can lead to quality loss
in sensitive fruits or vegetables (Wills et al., 1999, 2000). It is clear that this
molecule has a huge impact in conventional handling and distribution sys-
tems, and increased understanding of its biology and mechanisms of effect
on quality loss is essential to reducing loss during postharvest handling.
Ethylene is produced via the methionine (Yang) cycle (Figure 2.1)
in the cell wall–cell membrane complex and is only functional when the

Methylthioribose kinase 1-phosphate

butyric acid
Methylthioadenosin Methionine
nucleosidase Transaminase
5'-Methylthioadenosin Methionine

Fruit ripening, auxin, wounding or ⊕ S-Adenosyl-

SAM synthetase
infection, chilling, water stress, anoxia ACC Synthase L-methionine
id a

1-Amino-cyclopropane- CO2

N-Malonyl-ACC (MACC) carboxylate Ethylene (C2H4)

yl- (ACC) −
on ⊕ −
al 1-M
N se
len CO

C fera
Low emperat rs
high l scaveng

A an s
thy high

tr Ethylene receptor


ou ing,

en, c re, free

gen en
exo it rip


Figure 2.1  Representation of the methionine (Yang) cycle showing the

control points in the cycle in regard to ethylene production. (Adapted from
Wang, K.L.-C. et al., The Plant Cell (Suppl.), S131–S151, 2002; Blankenship,
S.M., and Dole, J.M., Postharvest Biology and Technology 28, 1–25, 2003;
Saltveit, M.E., Postharvest Biology and Technology 15, 279–292, 1999.)

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

membrane is associated with the cell wall (Mattoo and Lieberman, 1977).
This understanding of the localization of production may explain the
observations that ethylene production is transient during ripening and
senescence or in response to wounding (Figure 2.2). The latter stages of
senescence involve cell wall breakdown and cell membrane disintegration
(Hurkman and Kennedy, 1975), which would imply that the association
of membrane with cell wall is lost. Also in wounding, cell membranes are
known to lose their functionality (Toivonen and DeEll, 2002), again imply-
ing the loss of cell wall membrane associations that exist in intact cells.
Transient ethylene production occurs only at the onset of climacteric ripen-
ing and senescence (Biale et al., 1954; Aharoni and Lieberman, 1979) or
early in the wound response (Toivonen and DeEll, 2002), which is consis-
tent with this hypothesis.
Fruits and vegetables have varying sensitivities to ethylene, and
responses to ethylene are manifested in many different ways. Table 2.1
shows the relative sensitivity of many fruits and vegetables commonly mar-
keted in North America. It is also important to note that many of the sensi-
tive fruits and vegetables can also produce significant amounts of ethylene,
because ethylene production is linked to the ripening of that fruit. Table 2.2
lists the responses that are evoked with exposure to ethylene by various

Climacteric peak
Carbon dioxide production



Non-climacteric pattern


Figure 2.2  Respiratory patterns for climacteric and nonclimacteric fruit.

(From Saltveit, M.E., in Gross, K.C. et al. (eds.), The Commercial Storage
of Fruits, Vegetables, and Florist and Nursery, USDA Handbook 66, U.S.
Department of Agriculture, Agricultural Research Service, Beltsville, MD,


Table 2.1  Relative Production Rates and Sensitivity to Ethylene in Selected

Fruits and Vegetables
Relative Ethylene Relative Sensitivity to
Fruit or Vegetable Production Rate Ethylene
Apple Very high High
Globe artichoke Very low Low
Asparagus Very low Moderate
Avocado High High
Banana Moderate Moderate
Green bean Low Moderate
Raspberry Low Low
Strawberry Low Low
Broccoli Very low High
Cabbage Very low Moderate to high
Carrots Very low High
Cauliflower Very low High
Celery Very Low Moderate
Cherries Very low Low
Cilantro Very low High
Citrus fruit Very low Moderate
Corn Very low Low
Cucumber Low High
Eggplant Low Moderate
Garlic Very low Low
Grape Very low Low
Chives Low Moderate
Kiwifruit Low High
Leafy greens Very low High
Lettuce Very low High
Mango Moderate Moderate
Mushrooms Very low Moderate
Mature onions Very low Low
Nectarines, plums, and peaches Moderate Moderate
European pear High High
Peppers Low Low
Pineapple Low Low
Potato Very low Medium
Squash Low Medium

(Continued )

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

Table 2.1  Relative Production Rates and Sensitivity to Ethylene in Selected

Fruits and Vegetables
Relative Ethylene Relative Sensitivity to
Fruit or Vegetable Production Rate Ethylene
Sweet potato and yam Very low Low
Tomato, mature green Very low High
Tomato, firm ripe High Low
Watermelon Very low High
Source: Extracted from Cantwell, M., in Kader, A.A. (ed.), Postharvest Technology
of Horticultural Crops, Agriculture and Natural Resources Publication
3311, University of California, Oakland, 2002, pp. 511–518.

Table 2.2  Changes or Disorders Caused by Ethylene Exposure for Selected

Fruits and Vegetables
Fruits or Vegetables Affected Response to Ethylene Exposure
Apple, avocado, pear, stone Accelerated climacteric ripening, softening
fruits, mature green tomato,
melons, watermelon, banana
Carrot, parsnip, celery Production of phytoalexins (iso- or
furanocoumarins) leading to bitterness;
furanocoumarins can cause contact
dermatitis on exposure to UV light when
handling vegetables with high contents
Leafy vegetables, cruciferous top Premature yellowing and senescence
vegetables, green asparagus,
herbs, citrus, cucumbers
Iceberg lettuce Postharvest foliar russet spotting
Asparagus Toughening through induction of lignification
Potatoes, bulb onions Sprouting at high concentrations
Potatoes Inhibition of sprouting at low concentrations
Mushrooms Browning of caps
Sweet potatoes Hard core (inedible hardening of flesh
evidenced upon cooking)
Sources: Information compiled from Timbie, M., and Haard, N.F. et al., Journal
of Food Science 42, 491–493, 1977; Gross, K.C. et al., The Commercial
Storage of Fruits, Vegetables, and Florist and Nursery Stocks, USDA
Handbook 66, U.S. Department of Agriculture, Agricultural Research Service,
Beltsville, MD, 2004,; Toivonen,
P.M.A., in Hui, Y.H. et al. (eds.), Handbook of Vegetables and Vegetable
Processing, Wiley-Blackwell Publishing, Ames, IA, 2010, pp. 199–220.


groups of fruits and vegetables. A part of produce incompatibility in stor-

age is based on the impact of ethylene on fruits and vegetables that do not
normally produce significant levels of ethylene but are sensitive to ethylene

2.2.2  Respiratory Characteristics:

Climacteric and Nonclimacteric
The respiratory behavior of fruit during ripening can be classified as
­climacteric or nonclimacteric in nature. In nonclimacteric fruit, the res-
piration rate declines slowly after harvesting. This respiratory decline is
generally assumed to occur as a consequence of depletion in substrates
available for respiration. In contrast, climacteric fruit exhibit rapid and dra-
matic respiratory increase during ripening (Figure 2.2). The climacteric
rise has been characterized as having four distinct phases, the first being
a transient decline in respiration rate at full fruit maturity, followed by a
rapid rise that reaches a maximum and then declines as the fruit tissue
deteriorates. In general, the climacteric peak is normally many orders of
magnitude above basal preclimacteric respiration.
The respiratory behavior has implications for the storage and
­handling of fruits or fruit-vegetables. In general, nonclimacteric fruits or
fruit-vegetables do not exhibit accelerated ripening in response to ethyl-
ene exposure; however, they may show other ethylene responses, such as
­yellowing, secondary development, or secondary compound accumulation
(Saltveit, 1999).

2.2.3  Developmental or Maturity Stage at Harvest

Harvest maturity is an important subject, and much effort is placed in
selecting the optimal maturity stage for all fruits and vegetables to ensure
the best quality and storage potential. Watada et al. (1984) defined the
developmental stages for horticultural crops in great detail (Table 2.3).
The physiological implications of developmental stage and the degree of
maturity at that stage are tremendous, and a misunderstanding could lead
to poor decisions in postharvest handling protocols. For example, tomato
fruit must be at “mature green” when internally the seeds are tan in color
and there is a significant development of a gel in at least two locules of the
fruit (Sargent and Moretti, 2004). If the fruit has not reached this stage
of maturity at harvest, they will not ripen, even if exogenous ethylene is
applied (Sargent and Moretti, 2004). However, generally the identification
of mature green can be difficult in practice, and the commercial industry

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

Table 2.3  Classification of Fruits and Vegetables Based on Developmental

Phase and Anatomical Structure
Phase in Development Description of Fruit
Continuum or Vegetable Examples
Vegetative growth Sprouts Alfalfa, onion, watercress,
bean sprouts
Stems or leaves Asparagus, celery, spinach,
lettuce, cabbage, kohlrabi,
chard, endive, herbs
Inflorescences Artichokes, broccoli,

Late vegetative growth Partially developed Cucumber, green beans,

to physiological fruit okra, sweet corn, green
maturation peppers, summer squash

Ripening Fully developed fruit Apple, pear, citrus, tomato,

colored peppers, mango,
papaya, avocado, stone
fruit, melons, winter squash
Roots and tubers Carrot, onion, potato, garlic,
Jerusalem artichoke,
jicama, ginger
Source: Adapted from Watada, A.E. et al., HortScience 119, 20–21, 1984.

often harvests at a “breaker” stage to ensure that the fruit will respond to
ethylene and ripen after harvest.
The rate of developmental change is also important to understand in
handling fruits and vegetables. In general, those fruits and vegetables har-
vested at the vegetative growth stage to the physiological maturity stage
of the development continuum will have the most rapid metabolism and
hence shortest shelf potential (Toivonen, 2010). In addition, there is usu-
ally a higher respiratory rate, and it is well known that shelf life is consid-
ered to be inversely proportional to respiration rate (Saltveit, 2004). For
fruits and vegetables harvested at the ripening phase of the developmental
continuum, if the correct timing is chosen, respiration rates can be quite
low; for example, if potatoes are allowed to grow to maturity and are cured
before ­storage, they have much lower respiration rates than if they are
harvested as immature potatoes (Toivonen, 2010). Also, climacteric fruit,
which can exhibit high respiration rates in the latter stages of ripening,


when harvested at the early climacteric stage and properly stored, have
relatively low respiration rates and can be stored for long periods of time
(Saltveit, 2004; Watkins et al., 2004).
Because timing of harvest is an important factor in determining
storage life, various practical indices have been developed to aid in mak-
ing optimal harvest decisions, ranging from visual and tactile indices to
precise instrumental measures (Reid, 2002). The approach in each case
depends on monitoring visual, tactile, or compositional changes that occur
in the fruit or vegetable, leading to the horticultural optimal maturity for
­harvest. In the case of lettuce, this means assessing the solidity of the head;
in apples, this means measuring the loss of starch with iodine staining,
and in persimmons the tannin content (Reid, 2002). Clearly, each fruit or
vegetable has a distinct approach to assessing maturity for harvest. In addi-
tion, there continues to be challenges for fruits and vegetables or new culti-
vars emerging into the export markets, where practical and reliable harvest
maturity indices have yet to be developed.

2.2.4  Tolerance to Low Temperatures

Sensitivity to chilling-induced quality changes or tissue injuries is an
important issue for postharvest handling and storage of subtropical and
tropical fruits and vegetables, but it can also be of concern in the handling
and storage of temperate fruits and vegetables (Wang, 2004). The thresh-
old temperatures for chilling injury can range from 13°C down to 0°C;
however, the mechanism for chilling-related disorders involves changes
in membrane functionality (Wang, 1982). Secondary events stemming
from loss in membrane functionality lead to chilling-associated disor-
ders, which can include internal browning, soft scald, skin darkening or
discoloration, abnormal ripening or loss of ability to ripen, tissue necro-
sis, pitting, hardening, internal discolorations, loss of aroma production,
and decay (Figure 2.3). It should be noted that decay often is secondary
to physical injury induced by the chilling response (Morris, 1982).
There has been much research devoted to modulating or eliminating
chilling-induced injury (Wang, 1993). Many of the proposed approaches to
alleviating chilling injury involve mechanisms that cause the modification
of membrane lipid composition, allowing the membrane to sustain func-
tionality (Luengwilai et al., 2012). Other approaches involve modulating the
metabolism of the fruit or vegetable such that chilling-induced changes are
minimized (Wang and Qi, 1997). It is also known that the more mature a
fruit is, the less susceptible it is to chilling injury, presumably because more
mature fruit has a higher enzymatic or nonenzymatic antioxidant content,
which helps to protect membranes from oxidative injuries (Qian et al., 2013).

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

Chilling temperature

Chilling sensitive fruit

or vegetable

Primary response
Phase transition of membrane
from liquid crystalline to solid gel

Secondary responses
ROS accumulations
Ethylene production
Increased respiration
Decoupling of electron transport from
energy production
Alteration of cellular structure
Alterations in metabolic pathways

Short exposure
to pre-stressed
–processes reversible

Prolonged exposure

Symptoms of injury
Surface pitting
Internal breakdown
Loss of capacity to ripen
Aroma Loss

Figure 2.3  Schematic representation of primary and secondary c­hilling

responses and ensuing symptoms that develop should the exposure
be prolonged. (Adapted from Wang, C.Y., HortScience 17, 173–186,
1982; Sevillano, L. et al., Journal of the Science of Food and Agriculture 89,
555–573, 2009.)


More recently, efforts have been focused on directed breeding to

confer chilling resistance in new cultivars of fruits in particular (Martínez-
García et al., 2012; Dhanapal et al., 2012). While genetic markers for chill-
ing resistance are being developed, the trait is not straightforward, and
the linkage to specific gene regulation is not well understood at this time
(Martínez-García et al., 2012). This problem is likely due to the fact that,
physiologically, the expression of chilling symptoms used to develop genetic
markers may involve secondary processes (Wang, 1982), and so a clear rela-
tionship between chilling symptoms and the primary processes leading to
them is not easily defined. Nevertheless, continued efforts to use genetic
markers to increase chilling resistance is a worthwhile effort that will
allow identification of chilling-resistant germplasm from strategic crosses
between resistant and commercially desirable parents. Because of the com-
plex nature of the gene expression relationship to chilling symptoms, it
is likely that genetic engineering will not provide significant efficiency at
achieving commercially desirable fruit with chilling tolerance (Vinocur and
Altman, 2005).

2.3  Factors Affecting Physiology of Fruits

and Vegetables in Postharvest Systems

2.3.1 Temperature
Temperature influences the rate of all metabolic processes, including res-
piration. Metabolic processes all rely on enzymatic activity, which can be
further generalized as a chemical reaction. As such, effects of temperature
can be described by an Arrhenius plot, which is derived as
An enzymatic reaction can be generalized as

Substrate A + Substrate B → Product A + Product B

The rate of the reaction is defined as

V = k × [Substrate A] × [Substrate B]

where V is the velocity of the reaction and k is the velocity constant.

The constant k is determined from an Arrhenius plot, where

k = Ae RT

where A is a constant, µ is the activation energy, R is the perfect gas
­constant, and T is the absolute temperature.

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

The effect of temperature change on reaction rate is defined as the

quotient of the rate of change over a 10°C change in temperature (i.e., Q10).
The Q10 for a particular reaction or enzyme activity is calculated as

Vt +10
Q10 =

where Vt is the rate measured at a temperature (t) measured in degrees
Celsius and Vt+10 is the rate at 10°C higher than t.
The Q10 has been widely used in the context of respiratory behavior
cold storage and to explain the effect of low temperatures on enhancing
the storage life of produce (Nunes and Emond, 2003; Bron et al., 2005).
However, it has to be noted that not all enzymes involved in respiratory
and nonrespiratory pathways have identical Q10 values in the tempera-
ture ranges studied (0°C–20°C). Pollock and ap Rees (1975a) demon-
strated that enzymes involved with sugar metabolism have wide ranges in
Q10 ­values, between 2°C and 10°C, and this consequently explained some
of the changes associated with low temperature sweetening in potatoes
(Pollock and ap Rees, 1975b). It has also been shown that enzyme systems
can become acclimatized to low temperatures, becoming less sensitive
to temperature change over time, and thus having lower Q10 values after
time in storage (Atkin and Tjoelker, 2003). Rhodes et al. (1981) demon-
strated differential changes in enzymes involved with phenolic metabo-
lism in response to temperature and differential temporal changes in
these enzyme activities with time at low temperature storage. This work
clearly emphasizes the importance of temperature for imposing shifts in
metabolic pathways, and doing so in such a manner that relative contents
of phenolic constituents are modified, and this relationship also shifts
over time as some of the enzymes acclimate, and others do not, to low
While Q10 principles have been applied to enzymatic processes in
fruits and vegetables, temperature also has effects on physical move-
ments into, out of, and within fruit tissue; that is, rates of gaseous and
liquid diffusion in tissues can be influenced by temperature (Lammertyn
et al., 2001). This finding emphasizes how little is known about the com-
ponents of apparent Q10 values developed for postharvest analysis (Bron
et al., 2005).
The importance of low temperatures to controlling quality decline in
fruits and vegetables is well known (Nunes and Emond, 2003). However, a
scan of the literature indicates that there are limited reports on differen-
tial effects of low temperature on fruit and vegetable synthetic, catabolic,
and respiratory pathways. The lack of good tools to measure a broad range
of proteins (enzymes) and metabolic substrates and products has likely
­limited work in this area. However, with new tools and data processing


capability for proteomic and metabolomic analysis, the investigation of dif-

ferential changes in a broad range of enzymes and metabolites in metabolic
pathways has now become a reality (Pedreschi et al., 2009; Rudell et  al.,
2009). Hopefully this will lead to a better understanding of the effects of
storage regimes, including temperature, on respiratory, metabolic, and
quality changes in fruits and vegetables.

2.3.2 Humidity
Humidity is the second most important factor, after temperature, for mod-
ulating changes in physiology and the quality of fruits and vegetables in
postharvest systems. The humidity of the air surrounding a fruit or veg-
etable exerts some vapor demand on the produce if the humidity is below
100%, which is normally the case in postharvest systems. Water loss can be
­calculated as

(P − P ) × A
i a f

( RD × T ) × r

where J is the water loss in weight of water per unit time per surface area
of the fruit, Pi and Pa are the steady-state vapor pressures of water in the
intercellular spaces of the fruit and in the atmosphere, respectively, A f is
the surface area of the fruit, R D is the gas constant, T is the absolute tem-
perature in degrees Kelvin, and r is the specific resistance of the fruit to
water loss. While this model provides precise estimates of water loss, it is
essential to determine the surface area of the fruit and the specific resis-
tance of that fruit to water loss. In practice, it is more important to get an
estimate of the Pi − Pa component of the water loss model so that relative
changes in driving force for water loss (i.e., vapor pressure deficit) can
be quantified, allowing evaluation of the effect of changes in postharvest
handling protocols on relative water loss from the fruit.
In order to calculate vapor pressure deficit, the saturation vapor pres-
sure of the fruit or vegetable and the surrounding air must be calculated.
Saturation vapor pressure (VPs ) can be calculated as
17.27 × T
VPs = 0.6108 × e T + 237.3

where T is the temperature of the produce or the surrounding air in degrees
The intercellular spaces of the fruit or vegetable are considered to be
at saturation vapor pressure (Ben-Yehoshua and Rodov, 2003). However,
the air vapor pressure (VPa ) can be calculated as

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

RH a
VPa = × VPsa

where RHa is the relative humidity of the air in percent and VPsa is the calcu-
lated saturation vapor pressure of the air at that temperature.
Water vapor deficit (VPD) is the driving force for water loss from
fruits or vegetables and is calculated as

VPD = VPp − VPa

where VPp is the calculated saturation vapor pressure of the fruit or veg-
etable and VPa is the calculated vapor pressure of the surrounding air based
on its temperature and relative humidity.
It is a straightforward task to calculate vapor pressure deficits for
many postharvest handling scenarios to demonstrate the need for rapid pre-
cooling compared to room cooling. It can be demonstrated that warm fruit
will have higher integral vapor pressure deficits if room cooled, and hence
greater water loss than fruit that is rapidly forced-air cooled (Toivonen
and Hodges, 2011). Clearly, it is important to cool fruits and vegetables to
temperatures as close to that of the storage room or transport container as
soon as possible in order to minimize water loss. In addition, it is impor-
tant to manage the relative humidity in a storage room or container such
that it is maintained as high as is suitable for a particular fruit or vegetable
(Cantwell, 2002).
As implied earlier, there are a number of factors that determine the
resistance of produce to water loss—the composition and thickness of
the epidermis; the presence of surface structures on the epidermis, such
as hairs; the number or presence of stomata (or lenticels); and the mor-
phological structure (a bulky organ with a low surface area-to-volume
ratio vs. a leafy green with a high surface area-to-volume ratio). The
resistance to water loss can be modulated with wax coatings or applica-
tion of packaging to control humidity around the produce (Toivonen and
Hodges, 2011).
If water loss is not controlled, there is certainly a loss of turgor leading
to wilting, but there are also a few other metabolic changes that can occur
(Toivonen and Hodges, 2011). The enzyme activities of polygalacturonase
and pectinesterase increase, leading to loss of cell wall structure and con-
comitant increases in soluble sugars in tomato (Inari et al., 2002). Similar
results have been found with cucumbers, in which water stress caused an
upregulation of polygalacturonase activity, leading to the hypothesis that
the physical loss of water (i.e., loss of turgor in the tissue) was not the only
factor in causing softening of stressed fruit (Kubo et al., 2000). Water stress
also induces increased ethylene production (Kubo et al., 2000), and this


may explain accelerated ripening in water-stressed bananas (Burdon et al.,

1994) and accelerated senescence in bell peppers (Lurie et al., 1986).

2.3.3  Atmospheric Modification

Modified and controlled atmospheres have been long used to modulate
fruit and vegetable physiology and metabolism in order to extend their stor-
age lives (Kader and Saltveit, 2003a). A wide range of physiological effects
on fruits and vegetables in response to reduced O2 or elevated CO2 have
been documented (Kader and Saltveit, 2003a). While the prior review of
this subject area is complete and up to date, a few points are worth not-
ing in this discussion. In many cases, both reduced O2 and elevated CO2
result in inhibition of many pathways or enzyme activities; however, in
some cases (e.g., phenylalanine ammonia lyase [PAL] activity, succinic
acid and malic acid accumulation, and chilling injury) they may counteract
each other. Therefore, overall quality change may be difficult to manage
using controlled and modified atmospheres, since some pathways respond
­differently. In one case, it has been shown that a high CO2 atmosphere
improves overall quality retention of stored strawberries; however, malic
acid content losses increase and lead to potential flavor loss since malic acid
is an important aspect of fruit flavor (Holcroft and Kader, 1999).
While controlled and modified atmosphere protocols have been devel-
oped to optimize storage life of fruits and vegetables, there are risks of using
the technology in certain cases (Kader and Saltveit, 2003). These cases
include aggravation of disorders such as blackheart in potatoes, irregular
ripening of melons and tomatoes (when O2 < 2% and CO2 > 5%), develop-
ment of off-flavors (when O2 < 0.5% and CO2 > 20%), increased susceptibil-
ity to decay in produce that has been injured in extreme atmospheres, and
simulation of sprouting and inhibition of periderm development in root and
tuber vegetables. Therefore, it is important to understand the physiologi-
cal parameters of each fruit and vegetable and the risk–benefit relationship
for atmosphere modification. Atmosphere recommendations for most fruits
and vegetables are listed by Cantwell (2002).
An emerging area of interest is the use of dynamically controlled
atmosphere (DCA) storage, which in principle works through adjusting
atmospheres during storage to meet changes in fruit acclimation to stor-
age conditions (Kader and Saltveit, 2003a). The challenge for this new
approach is to have monitoring instrumentation that can sense fine physi-
ological changes in the fruit, enabling appropriate atmosphere adjustment
while avoiding tissue injury (Saltveit, 2003). It also must be noted that this
technology, like most, may not be beneficial for all cultivars, and there
are significant new considerations that must be met before successful

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

implementation can take place in commercial practice (Watkins, 2008).

There is much need for further research to better understand the effect
of dynamically controlled ultralow oxygen on fruit physiology and

2.3.4  Abiotic Stresses

Postharvest handling by definition imposes abiotic stress on a fruit or
vegetable (Toivonen and Hodges, 2011). However, the fruit or vegetable
may be subject to preharvest factors or stresses that also lead to modified
stress susceptibility in postharvest handling and storage (Toivonen, 2003;
Toivonen and Hodges, 2011). In general, three possible scenarios can exist
when a fruit or vegetable has been harvested: (1) it can be in an unstressed
state and, as such, can withstand many stresses imposed by postharvest
processes; (2) it can have been subjected to mild stresses, which enhance
its resistance to stresses imposed by postharvest processes; or (3) it can
have been subjected to severe stresses, which weaken its resistance to sub-
sequent stress exposure in postharvest processes.
Ripeness stage, storage temperature, ethylene, and oxidative stress
all affect peel phytosterol metabolism in apples and thus influence the
development of superficial scald (Rudell et al., 2011). There has been some
work showing that sun exposure on the tree may lead to invisible dam-
age on apple fruit (Kuckenberg et al., 2008), which may contribute to the
de­velopment of postharvest scald (Hernandez et al., 2014). These observa-
tions suggest that scald is associated with chilling responses. When review-
ing the approaches being adopted to control disorders such as superficial
scald in apples (Rudell et al., 2011), it becomes clear that these are the same
approaches that have been adopted to modulate chilling injury in other
fruits and vegetables (Sevillano et al., 2009).
Mechanical injury is a consequence of physical handling procedures
employed during the harvest and packing of fruit. If often leads to bruis-
ing in fruit such as apple, and the visual appearance of the bruising can be
reduced by an “apparent recovery” or suberization period after the bruising
injury is incurred (Toivonen et al., 2007). Proteomic analysis of response
to mechanical injury in apples has shown that response to stress proteins,
many of which are considered to be pathogen resistance–related proteins,
was upregulated with wounding (Buron-Moles et al., 2014). This supports
the overall hypothesis of cross-tolerance mechanisms in fruit and vegeta-
ble tissues, where it is suggested that the responses to biotic and abiotic
stresses share many common gene and metabolome expression pathways
(Toivonen, 2004).


2.4  Physiological Changes Occurring during

Postharvest Handling or Storage

2.4.1  Depletion of Respiratory Substrate

Despite attempts to inhibit respiration in postharvest systems with the
use of low temperatures, controlled or modified atmospheres (Kader and
Saltveit, 2003b), and other technologies, such as 1-methylcyclopropene
(Blankenship and Dole, 2003), there is an inevitable loss of respiratory
substrate over time, which can be significant with long storage durations
extending several months. The consequence of such depletion is clearly
associated with important quality attributes of fruits and vegetables.
Loss of malic acid content is a significant problem in the long-term stor-
age of some fruits, such as apple, where malic acid is an important aspect
of sensory quality (Jan and Rab, 2012). While the sugar concentration
of apples generally increases during long-term storage, the dry matter
content declines (Jan and Rab, 2012). Dry matter has also been recently
identified as an indicator of poststorage quality in apples (Palmer et al.,
2010). It is clear from these observations that dry matter and respiratory
organic acids such as malic acid are depleted due to respiratory activity
over long-term, low-temperature storage, whereas sugar concentrations
are maintained or increased. It may be that low-temperature effects on
sugar metabolic enzymes (ap Rees et al., 1981) are partially responsible
for this apparently contradictory response. Dry matter content at harvest
has also been shown to be important for the short-term storage of broccoli
microgreens (Kou et al., 2014), suggesting that the relationship between
dry matter and sustained respiration during storage holds true for fruits
and vegetables.
The potential storage life of fruits and vegetables is partially deter-
mined by their relative respiration rates; however, the other side of the issue
is that storage life comes to an end when a critical threshold for a respira-
tory substrate has been reached (Kader and Saltveit, 2003b). Respiration
is essential for the living tissues to maintain cellular organization and
function through the maintenance of adequate adenosine triphosphate
(ATP) titers in the cells, allowing the ongoing basal metabolic activities
to continue. As respiratory substrates are depleted, the ability to regener-
ate ATP declines, which could potentially affect fruit tissues and quality
retention. There has been some work done to demonstrate the importance
of the energy state on the postharvest development of quality defects in
apple and pear fruit (Saquet et al., 2000). More recently, it has been sug-
gested that regulation of the energy state in fruit may be the key to manag-
ing ripening and senescence in storage (Wang et al., 2013). The concept

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

of energy homeostasis has emerged from this work (Figure  2.4), which
is congruent with earlier understanding in regard to the maintenance of
cellular organization and function. Clearly there is significant understand-
ing yet to be had in regard to the maintenance metabolism of tissues in

Stress (senescence)

+ −

ABA ATP Deficit

SnRK Signaling

ATP Transporting ATP Synthesising ATP Dissipating Energy homeostasis


Fruit Quality

Figure 2.4  Possible mechanism to account for energy regulation in senescent

litchi fruit. An energy deficit in intensive respiration during the senescence pro-
cess may be sensed by sucrose non-fermenting-1-related kinase (SnRK), which
controls the expression and phosphorylation of key metabolic enzymes, and
this might involve ATP synthase, ADP/ATP carrier (AAC), alternative oxidase
(AOX), and uncoupling mitochondrial protein (UCP). The energy homeostatic
condition can be maintained to a certain extent, but the equilibrium is ulti-
mately disrupted, which is correlated with fruit deterioration. (From Wang, H.
et al., BMC Plant Biology 13, 55, 2013.)


2.4.2  Hormonal Effects

There are two aspects of hormonally associated changes in postharvest
systems. The first is related to endogenous shifts in hormone levels and
the consequent metabolic changes, and the second involves the effects of
exogenously applied hormones. Early work focused on measures of endog-
enous hormones and developmental changes in fruit and vegetable tissues
(Ludford, 2003). Work on exogenously applied phytohormones followed
that research (Ludford, 2003; Klein and Goldshmidt, 2005) and continues
to present. However, only recently have advances in this area occurred on
a broader genomic, proteomic, and metabolomic scale. Genomic studies
evaluating the response of fruit to endogenous hormone contents have pro-
vided a broader, more integrated understanding of response mechanisms
(Tacken et al., 2010; Devoghalaere et al., 2012; Matsuo et al., 2012). This
broader and more integrated analysis of responses is already better defin-
ing the complex metabolic changes that are induced (Pech et al., 2008),
and this is expected to lead to a more useful and reliable understanding of
control mechanisms.
Ethylene is probably the most studied hormone in postharvest research
literature. It is associated with the onset of ripening and senescence (refer
to discussion in Section 2.2.1), and as such, it has been employed to acceler-
ate or initiate ripening in fruits that are harvested green. Ethylene is rou-
tinely used for ripening green bananas and tomatoes, degreening lemons,
and conditioning pears that are harvested green (Toivonen, 2010; Sugar and
Basile, 2013). It has been discovered that higher levels of ethylene expo-
sure can be applied to inhibit sprouting in potato tubers (Daniels-Lake
et al., 2005). While ripening in climacteric fruit is considered to be highly
associated with ethylene, it is clear that changes at the molecular level are
­complex, involving both the gene transcription level (Nishiyama et  al.,
2007) and the posttranscriptional level, where micro-RNAs may be oper-
ating to modulate gene expression through differential silencing of target
genes involved with ethylene biosynthesis and signaling (Zuo et al., 2012).
Also, it is becoming clear that there is ethylene-dependent and -­independent
regulation of metabolic pathways associated with ripening (Zuo et al., 2012),
the impact of which is yet to be fully understood.
Gibberellic acid (GA) has been used for preharvest management of
fruit quality in many fruits. It can increase citrus size and reduce seed set in
facultatively parthenocarpic citrus cultivars (Iglesias et al., 2007). In sweet
cherries, GA has been adopted widely in commercial practice to delay
fruit maturation and increase fruit size and firmness (Canli and Orhan,
2009). Similar responses have been found in seedless table grapes with
GA applied at times soon after fruit set (Singh et al., 1978). The basis for
these responses is thought to be that endogenous GA normally produced

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

by the seed is either replaced or enhanced with the exogenous application.

Exogenous GA application has been shown to induce increases in fruit
length and girth, as well as soluble solids contents (Banerjee and Basu,
1992). GA has also been shown to downregulate polygalacturonase in sweet
cherries, which may provide some insight into its effect on fruit firmness
(Choi et al., 2002). Other work, with persimmon, has shown that exogenous
GA application results in modulation of expression for an expansin gene
(Zhang et al., 2012). Expansin is associated with fruit size or advance in
ripening of fruit (Brummell et al., 1999; Hiwasa et al., 2003), which may
explain the increased fruit size and delay in ripening of sweet cherries
treated with exogenous GA (Canli and Orhan, 2009). Also, higher levels of
production of endogenous GA early in development in Japanese pears have
been shown to relate to larger fruit size in one cultivar when compared to
one that produces very low levels (Zhang et al., 2007). In relation to matu-
rity, exogenous GA application has been shown to reduce sugar supply to
orange peel, suggesting that a delay of peel maturation could be directly
due to limitation in sugar substrate to that tissue (Fidelibus et al., 2008).
Clearly, a more in-depth and integrated analysis of GA effects on gene
expression and metabolome is necessary to better understand the varied
physiological effects of this hormone, especially since it is used extensively
in the commercial horticulture to modulate fruit quality.
Abscisic acid (ABA) has been shown to have a range of effects when
applied to different fruits. In southern highbush blueberry, ABA application
resulted in delayed maturity and firmer fruit, but had no other quality or
nutritional effects (Buran et al., 2012). In contrast, exogenous ABA enhanced
antioxidant capacities, anthocyanins, and flavonol contents in muscadine
grapes (Sandhu et al., 2011), although not all cultivars tested showed a sig-
nificant response. Cantín et al. (2007) found that exogenous ABA treatment
improved color development and visual appearance of Crimson seedless
grapes in comparison to industry existing practices. Jiang and Joyce (2003)
found that exogenous ABA accelerated softening and color development in
strawberries, and this was linked directly to effects of ABA on ethylene syn-
thesis and phenylalanine ammonia lyase upregulation, respectively. High
levels of ABA production have been linked to small fruit size in Japanese
pear (Zhang et al., 2007). Sun et al. (2013) recently investigated gene expres-
sion in melon fruit development and ripening, finding that genes encoding
non-ethylene-dependent ripening changes were regulated by ABA, and this
includes the induction of climacteric ethylene production. They suggested
that ABA plays a significant role in the early stages of ripening in the fruit,
and that ethylene takes over the major regulatory role at later stages.
Exogenous auxin application has been shown to increase fruit size,
enhance color, and advance ripening in the Maxim® apricot (Bregoli
et al., 2010). In a more critical study, a large number of genes have been


shown to be either repressed or induced by exogenous auxin application in

­strawberry (Aharoni et al., 2002). In general, the auxin-related changes in
gene expression were linked to cell wall metabolism and stress response
(including ethylene production). In a more detailed study, auxin response
was found to correlate to a quantitative trait loci (QTL) associated with fruit
weight in apples (Devoghalaere et al., 2012). Their results also suggest that
exogenous application of auxin at low concentrations (10 −7 M indoleacetic
acid) in early fruit development could enhance apple diameter; however,
higher concentrations (10 −5 M indoleacetic acid or higher) could negate
the size increase response or even reduce fruit size relative to untreated
controls. Response to auxin is very much determined by the timing of exog-
enous application and the concentration that is applied. However, in the end,
it seems that modulating auxin response is best managed through QTL
analysis and directed breeding approaches since factors affecting exog-
enous application may not be easily managed in a wide range of cultivars,
and the auxin response is extremely complex.
Exogenous cytokinin applications have been shown to delay loss of
chlorophyll and senescence in green leafy vegetables (Ludford et al., 2003).
This response has been linked to upregulation of genes for light-­harvesting
cytokinin-binding proteins and stimulation of chloroplast-encoded tran-
scription in vegetative tissues (Kulaeva et al., 2000; Zubo et al., 2008).
However, more recently it has been demonstrated that increases in endog-
enous cytokinin levels in kiwifruit lead to effects on response regulators
to the STAY-GREEN2 gene (Pilkington et al., 2013). This work suggests
that cytokinin is important to chlorophyll contents of green developing
­k iwifruit; however, it does not show that cytokinin is responsible in control-
ling the ripening-associated loss of chlorophyll in the fruit. In tomato fruit,
cytokinin has been demonstrated to promote the activity of genes involved
with cell division during early fruit development (Matsuo et al., 2012).
Phytohormones exert profound regulatory control over fruit and
vegetable development and final quality. The preceding discussion has
shown that the application of recent advances in metabolomic, proteomic,
and genomic analyses has created a greater understanding in hormone
effects on different aspects of fruit growth, development, and ripening.
However, endogenous phytohormones coexist in the intact fruit and vary
in level temporally during fruit development and ripening, and may modu-
late or even counteract each other’s activities in some cases. For example,
auxin and abscisic acid behave antagonistically in the expression of a
SHAT TERPROOF-like gene in the ripening of strawberry (Daminato et al.,
2013). The SHAT TERPROOF-like gene has a profound effect on modulat-
ing the expression of many genes regulating the ripening of the strawberry.
In another example, it has been demonstrated that exogenous cytokinin-
induced parthenocarpy in tomato was partially due to the effects on cyto-
kinin in modulating GA and auxin metabolism (Ding et al., 2013). This last

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

report highlights the importance of the coordinated regulatory actions of

all phytohormones in effecting the development, growth, and ripening of
fruit. The understanding at the genome and metabolome level is only now
beginning to develop and in the future should explain the physiological
effects that have been documented in earlier literature.

2.4.3  Membrane Alterations

As mentioned above in the discussion surrounding chilling stress, altera-
tion of membrane function is the primary response to that stress. Chilling
and other stresses alter the function of membrane proteins and synthetic
pathways, leading to many changes in metabolite accumulations, includ-
ing o­ xygen free radicals (Alcázar et al., 2010). Consequent tertiary effects
include significant alterations in gene expression and loss of membrane
permeability and transport functions. Production of polyamines, which
is closely associated metabolically with ethylene synthesis (Alcázar
et al., 2010), has been shown to modify lipid bilayer membrane structure
and function such that increased tolerance to stress is realized. Other work
with apples has shown that storage stress leads to significant changes
in peel phytosterol metabolism (Rudell et al., 2011). Phytosterols are
­important to  membrane fluidity and function. Rudell et al. (2011) found
that steryl-6-O-fattyacyl β-D-glycoside accumulation in apple peel was an
indicator that the apples had suffered stress that could lead to superficial
scald injury of the peel. Others have observed oxidative injury membranes
in association with storage disorder development (Whitaker, 2004).

2.5  Conclusions and Future Perspectives

The physiology of harvested fruits and vegetables is very sensitive to condi-
tions of handling, and their sensitivity to handling can be modified by treat-
ments prior to harvest and after harvest. In the end, physiological sensitivity
and response are governed by gene, proteome, and metabolome expression
in the fruit or vegetable tissues. The research in postharvest physiology has
enjoyed many decades of advances using limited analysis techniques and rela-
tively easily measured parameters to judge physiological change in response
to handling and storage conditions. However, as new data are emerging from
talented research groups and networks, it is becoming clear that nuances
and unexpected outcomes noted in prior postharvest studies can potentially
be better understood by integrated use of emerging techniques in genom-
ics, proteomics, and metabolomics. The challenge of such approaches is that
they are quite costly and are more realistically supported through leveraged
funding and effort from a number of institutions and countries. Research


groups providing the most comprehensive results are those who have truly
multidisciplinary collaborators, including crop production physiologists,
postharvest physiologists, chemists, and molecular b ­ iologists. This new
era of postharvest physiology requires new ways of working together and a
multidisciplinary approach to provide both perspective on the physiological
problem and a detailed understanding of the ­molecular mechanisms that
will allow us to better predict postharvest quality outcomes from treatment
applications and for directed breeding programs.

Aharoni, N., and Lieberman, M. 1979. Patterns of ethylene production in
senescing leaves. Plant Physiology 64, 796–800.
Aharoni, A., Keizer, L.C.P, Van Den Broeck, H.C., Blanco-Portales, R., Muñoz-
Blanco, J., Bois, G., Smit, P., De Vos, R.C.H., and O’Connell, A.P. 2002.
Novel insight into vascular, stress, and auxin-dependent and -independent
gene expression programs in strawberry, a non-climacteric fruit. Plant
Physiology 129, 1019–1031.
Alcázar, R., Altabella, T., Marco, F., Bortolotti, C., Reymond, M., Koncz, C.,
Carrasco, P., and Tiburcio, A.F. 2010. Polyamines: Molecules with regu-
latory functions in plant abiotic stress tolerance. Planta 231, 1237–1249.
ap Rees, T., Dixon, W.L., Pollock, C.J., and Franks, F. 1981. Low temperature
sweetening of higher plants. In Friend, J., and Rhodes, M.J.C. (eds.),
Recent Advances in the Biochemistry of Fruits and Vegetables. Academic
Press, London, pp. 41–61.
Atkin, O.K., and Tjoelker, M.G. 2003. Thermal acclimation and the dynamic
response of plant respiration to temperature. Trends in Plant Science 8,
Banerjee, S., and Basu, P.S. 1992. Hormonal regulation of flowering and fruit
development: Effect of gibberellic acid and ethrel on fruit setting and
development of Momordica charantia L. Biologia Plantarum 34, 63–70.
Ben-Yehoshua, S., and Rodov, V. 2003. Transpiration and water stress. In
Bartz, J.A., and Brecht, J.K. (eds.), Postharvest Physiology and Pathology
of Vegetables, 2nd ed. Marcel Dekker, New York, pp. 111–159.
Biale, J.B., Young, R.E., and Olmstead, A.J. 1954. Fruit respiration and ethyl-
ene production. Plant Physiology 29, 168–174.
Blankenship, S.M., and Dole, J.M. 2003. 1-Methylcyclopropene: A review.
Postharvest Biology and Technology 28, 1–25.
Bregoli, A.M., Fabbroni, C., Raimondi, V., and Costa, G. 2010. Improving
colour and size of apricot fruit by means of exogenous auxin applica-
tion. Acta Horticulturae 862, 365–372.
Bron, I.U., Ribeiro, R.V., Cavalini, F.C., Jacomino, A.P., and Trevisan, M.J.
2005. Temperature-related changes in respiration and Q10 coefficient of
guava. Scientia Agricola 62, 458–463.

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

Brummell, D.A., Harpster, M.H., and Dunsmuir, P. 1999. Differential

­expression of expansion gene family members during growth and ripen-
ing of tomato fruit. Plant Molecular Biology 39, 161–191.
Buran, T.J., Sandhu, A.K., Azeredo, A.M., Bent, A.H., Williamson, J.G., and
Gu, L. 2012. Effects of exogenous abscisic acid on fruit quality, anti-
oxidant capacities, and phytochemical contents of southern high bush
blueberries. Food Chemistry 132, 1375–1381.
Burdon, J.N., Dori, S., Lomaniec, E., Marinansky, R., and Pesis, E. 1994.
The postharvest ripening of water stressed banana fruits. Journal of
Horticulture Science 69, 799–804.
Buron-Moles, G., Torres, R., Amoako-Andoh, F., Viñas, I., Teixidó, N., Usall,
J., Keulemans, W., and Davey, M.W. 2014. Analysis of changes in pro-
tein abundance after wounding in ‘Golden Delicious’ apples. Postharvest
Biology and Technology 87, 51–60.
Canli, F.A., and Orhan, H. 2009. Effects of preharvest gibberellic acid applica-
tions on fruit quality of ‘0900 Ziraat’ sweet cherry. HortTechnology 19,
Cantín, C.M., Fidelibus, M.W., and Crisosto, C.H. 2007. Application of abscisic
acid (ABA) at veraison advanced red color development and maintained
postharvest quality of ‘Crimson Seedless’ grapes. Postharvest Biology
and Technology 46, 237–241.
Cantwell, M. 2002. Summary table of optimal handling conditions for fresh
produce. In Kader, A.A. (ed.), Postharvest Technology of Horticultural
Crops. Agriculture and Natural Resources Publication 3311. University
of California, Oakland, pp. 511–518.
Choi, C., Wiersma, P.A., Toivonen, P., and Kappel, F. 2002. Fruit growth,
firmness and cell wall hydrolytic enzyme activity during development
of sweet cherry fruit treated with gibberellic acid (GA3). Journal of
Horticultural Science and Biotechnology 77, 615–621.
Daminato, M., Guzzo, F., and Casadoro, G. 2013. A SHATTERPROOF-like
gene controls ripening in non-climacteric strawberries, and auxin
and abscisic acid antagonistically affect its expression. Journal of
Experimental Botany 64, 3775–3786.
Daniels-Lake, B.J., Prange, R.K., Nowak, J., Asiedu, S.K., and Walsh, J.R. 2005.
Sprout development and processing quality changes in potato tubers
stored under ethylene. 1. Effects of ethylene concentration. American
Journal of Potato Research 82, 389–397.
Devoghalaere, F., Doucen, T., Guitton, B., Keeling, J., Payne, W., Ling, T.J.,
Ross, J.J., Hallett, I.C., Gunaseelan, K., Dayatilake, G.A., Diak, R., Breen,
K.C., Tustin, D.S., Costes, E., Chagné, D., Schaffer, R.J., and David,
K.M. 2012. A genomics approach to understanding the role of auxin in
apple (Malus × domestica) fruit size control. BMC Plant Biology 12, 7.
Dhanapal, A.P., Martínez-García, P.J., Gradziel, T.M., and Crisosto, C.H.
2012. First genetic linkage map of chilling injury susceptibility in peach
(Prunus persica (L.) Batsch) fruit with SSR and SNP markers. Journal of
Plant Science and Molecular Breeding, doi: 10.7243/2050-2389-1-3.


Ding, J., Chen, B., Xia, X., Mao, W., Shi, K., Zhou, Y., and Yu, J. 2013. Cytokinin-
induced parthenocarpic fruit development in tomato in partly depen-
dent on enhanced gibberellin and auxin biosynthesis. PLoS ONE  8,
e70080. doi: 10.1371/journal.pone.0070080.
Fidelibus, M.W., Koch, K.E., and Davies, F.S. 2008. Gibberellic acid alters
sucrose, hexoses, and their gradients in peel tissues during color
break delay in ‘Hamlin’ orange. Journal of the American Society for
Horticultural Science 133, 760–767.
Gross, K.C., Wang, C.Y., and Saltveit, M. 2004. The Commercial Storage of
Fruits, Vegetables, and Florist and Nursery Stocks. USDA Handbook 66.
U.S. Department of Agriculture, Agricultural Research Service,
Beltsville, MD. (accessed October 1,
Hernandez, O., Torres, C.A., Moya-León, M.A., Opazo, M.C., and Razmilic, I.
2014. Roles of the ascorbate-gluathione cycle, pigments and phenolics
in postharvest ‘sunscald’ development on ‘Granny Smith’ apples (Malus
domestica Borkh.). Postharvest Biology and Technology 87, 79–87.
Hiwasa, K., Rose, J.K.C., Nakano, R., Inaba, A., and Kubo, Y. 2003. Differential
expression of seven α-expansin genes during growth and ripening of
pear fruit. Physiologia Plantarum 117, 564–572.
Holcroft, D.M., and Kader, A.A. 1999. Controlled atmosphere-induced
changes in pH and organic acid metabolism may affect color of stored
strawberry fruit. Postharvest Biology and Technology 17, 19–32.
Hurkman, W.J., and Kennedy, G.S. 1975. Ultrastructural changes of chlo-
roplasts in aging tobacco leaves. Proceedings of Indiana Academy of
Sciences 85, 89–95.
Iglesias, D.J., Cercós, M., Colmenero-Flores, J.M., Naranjo, M.A., Ríos, G.,
Carrera, E., Ruiz-Rivero, O., Lliso, I., Morillon, R., Tadeo, F.R., and
Talon, M. 2007. Physiology of citrus fruiting. Brazilian Journal of Plant
Physiology 19, 333–362.
Inari, T., Yamauhi, R., Kato, K., and Takeuchi, T. 2002. Changes in pectic poly-
saccharides during the ripening of cherry tomato fruits. Food Science
and Technology Research 8, 55–58.
Jan, I., and Rab, A. 2012. Influence of storage duration on physic-­chemical
changes in fruit of apple cultivars. Journal of Animal and Plant
Science 22, 708–714.
Jiang, Y., and Joyce, D.C. 2003. ABA effects on ethylene production, PAL
activity, anthocyanin and phenolic contents of strawberry fruit. Plant
Growth Regulation 39, 171–174.
Kader, A.A., and Saltveit, M.E. 2003a. Atmosphere modification. In Bartz,
J.A., and Brecht, J.K. (eds.), Postharvest Physiology and Pathology of
Vegetables. Marcel Dekker, New York, pp. 229–246.
Kader, A.A., and Saltveit, M.E. 2003b. Respiration and gas exchange. In
Bartz, J.A., and Brecht, J.K. (eds.), Postharvest Physiology and Pathology
of Vegetables. Marcel Dekker, New York, pp. 7–29.

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

Klein, J.D., and Goldshmidt, E.E. 2005. Hormonal regulation of ripening

and senescence phenomena. In Ben-Yehoshua (ed.), Environmentally
Friendly Technologies for Agricultural Produce Quality. Taylor & Francis
Group, Boca Raton, FL, pp. 315–331.
Kou, L., Yang, T., Luo, Y., Liu, X., Huang, L., and Codling, E. 2014. Pre-harvest
calcium application increases biomass and delays senescence of broc-
coli microgreens. Postharvest Biology and Technology 87, 70–78.
Kubo, Y., Xue, Y.B., Nakatsuka, A., Mathooko, F.M., Inaba, A., and Nakamura,
R. 2000. Expression of a water stress-induced polygalacturonase
gene in harvested cucumber fruit. Journal of the Japanese Society for
Horticultural Science 69, 273–279.
Kuckenberg, J., Tarachnyk, I., and Noga, G. 2008. Evaluation of fluorescence
and remission techniques for monitoring changes in peel chlorophyll
and internal fruit characteristics in sunlight and shaded sides of apple
fruit during shelf life. Postharvest Biology and Technology 48, 231–241.
Kulaeva, O.N., Karavaiko, N.N., Selivankina, S.Y., Kusnetsov, V.V.,
Zemlyachenko, Y.V., Cherepneva, G.N., Maslova, G.G., Lukevich, T.V.,
Smith, A.R., and Hall, M.A. 2000. Nuclear and chloroplast cytokinin-
binding proteins from barley leaves participating in transcription regu-
lation. Plant Growth Regulation 32, 329–335.
Lammertyn, J., Franck, C., Verlinden, B.E., and Nicolaï, B.M. 2001.
Comparative study of O2, CO2 and temperature effect on respiration
between ‘Conference’ pear cell protoplasts in suspension and intact
pears. Journal of Experimental Botany 52, 1769–1777.
Ludford, P.M. 2003. Hormonal changes during postharvest. In Bartz, J.A., and
Brecht, J.K. (eds.), Postharvest Physiology and Pathology of Vegetables.
Marcel Dekker, New York, pp. 31–77.
Luengwilai, K., Beckles, D.M., and Saltveit, M.E. 2012. Chilling-injury of har-
vested tomato (Solanum lycopersicum L.) cv. Micro-Tom fruit is reduced
by temperature pre-treatments. Postharvest Biology and Technology 63,
Lurie, S., Shapiro, B., and Ben-Yehoshua, S. 1986. Effects of water stress and
degree of ripeness on rate of senescence of harvested bell pepper fruit.
Journal of the American Society for Horticultural Science 111, 880–885.
Martínez-García, P.J., Peace, C.P., Parfitt, D.E., Ogundiwin, E.A., Fresnedo-
Ramírez, J., Dandekar, A.M., Gradziel, T.M., and Crisosto, C.H. 2012.
Influence of year and genetic factors on chilling injury susceptibility in
peach (Prunus persica (L.) Batsch). Euphytica 185, 267–280.
Matsuo, S., Kikuchi, K., Fukuda, M., Honda, I., and Imanishi, S. 2012. Roles
and regulation of cytokinins in tomato fruit development. Journal of
Experimental Botany 63, 5569–5579.
Mattoo, A.K., and Lieberman, M. 1977. Localization of the ethylene-­
synthesizing system in apple tissue. Plant Physiology 60, 794–799.
Morris, L.L. 1982. Chilling injury of horticultural crops: An overview.
HortScience 17, 161–162.


Nishiyama, K., Guis, M., Rose, J.K.C., Kubo, Y., Bennett, K.A., Wangjin, L.,
Kato, K., Ushijima, K., Nakano, R., Inaba, A., Bouzayen, M., Latché, A.,
Pech, J.-C., and Bennett, A.B. 2007. Ethylene regulation of fruit softening
and cell wall disassembly in Charentais melon. Journal of Experimental
Botany 58, 1281–1290.
Nunes, M.C.N., and Emond, J.P. 2003. Storage temperature. In Bartz, J.A., and
Brecht, J.K. (eds.), Postharvest Physiology and Pathology of Vegetables.
Marcel Dekker, New York, pp. 209–228.
Palmer, J.W., Harker, F.R., Tustin, D.S., and Johnston, J. 2010. Fruit dry ­matter
concentration: A new quality metric for apples. Journal of the Science of
Food and Agriculture 90, 2586–2594.
Pech, J.C., Bouzayen, M., and Latché, A. 2008. Climacteric fruit ripening:
Ethylene-dependent and independent regulation of ripening pathways
in melon fruit. Plant Science 175, 114–120.
Pedreschi, R., Hertog, M., Robben, J., Lilley, K.S., Karp, N.A., Baggerman, G.,
Vanderleyden, J., and Nicolaï, B. 2009. Gel-based proteomics approach
to the study of metabolic changes in pear tissue during storage. Journal
of Agricultural and Food Chemistry 57, 6997–7004.
Pilkington, S.M., Montefiori, M., Galer, A.L., Emery, R.J.M., Allan, A.C., and
Jameson, P.E. 2013. Endogenous cyctokinin in developing kiwifruit is
implicated in maintaining fruit flesh chlorophyll levels. Annals of Botany
112, 57–68.
Pollock, C.J., and ap Rees, T. 1975a. Activities of enzymes of sugar metabolism
in cold-stored tubers of Solanum tuberosum. Phytochemistry 14, 613–617.
Pollock, C.J., and ap Rees, T. 1975b. Effect of cold on glucose metabolism by
callus and tubers of Solanum tuberosum. Phytochemistry 14, 1903–1906.
Qian, C., He, Z., Zhao, Y., Mi, H., Chen, X., and Mao, L. 2013. Maturity-
dependent chilling tolerance regulated by the antioxidative capacity in
postharvest cucumber (Cucumis sativus L.) fruits. Journal of the Science
of Food and Agriculture 93, 626–633.
Reid, M.S. 2002. Ethylene in postharvest technology. In Kader, A.A. (ed.),
Postharvest Technology of Horticultural Crops. Agriculture and Natural
Resources Publication 3311. University of California, Oakland,
pp. 149–162.
Rhodes, M.J.C., Wooltorton, L.S.C., and Hill, A.C. 1981. Changes in phenolic
metabolism in fruit and vegetable tissues under stress. In Friend, J., and
Rhodes, M.J.C. (eds.), Recent Advances in the Biochemistry of Fruits and
Vegetables. Academic Press, London, pp. 193–220.
Rudell, D.R., Buchanan, D.A., Leisso, R.S., Whitaker, B.D., Mattheis, J.P.,
Zhu, Y., and Varanasi, V. 2011. Ripening, storage temperature, ethylene
action, and oxidative stress alter apple peel phytosterol metabolism.
Phytochemistry 72, 1328–1340.
Rudell, D.R., Mattheis, J.P., and Hertog, M.L. 2009. Metabolomic change
precedes apple superficial scald symptoms. Journal of Agricultural and
Food Chemistry 57, 8459–8466.

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

Saltveit, M.E. 1999. Effect of ethylene on quality of fresh fruits and vegeta-
bles. Postharvest Biology and Technology 15, 279–292.
Saltveit, M.E. 2003. Is it possible to find an optimal controlled atmosphere?
Postharvest Biology and Technology 27, 3–13.
Saltveit, M.E. 2004. Respiratory metabolism. In Gross, K.C., Wang, C.Y.,
and Saltveit, M.E. (eds.), The Commercial Storage of Fruits, Vegetables,
and Florist and Nursery. USDA Handbook 66. U.S. Department of
Agriculture, Agricultural Research Service, Beltsville, MD. http://www. (accessed October 15, 2013).
Sandhu, A.K., Gray, D.J., Lu, J., and Gu, L. 2011. Effects of exogenous
abscisic acid on antioxidant capacities, anthocyanins, and flavonol con-
tents of muscadine grape (Vitis rotundifolia) skins. Food Chemistry 126,
Saquet, A.A., Streif, J., and Bangerth, F. 2000. Changes in ATP, ADP and
pyridine nucleotide levels related to the incidence of physiological dis-
orders in ‘Conference’ pears and ‘Jonagold’ apples during controlled
atmosphere storage. Journal of Horticultural Science and Biotechnology
75, 243–249.
Sargent, S.A., and Moretti, C.L. 2004. Tomato. In Gross, K.C., Wang, C.Y.,
and Saltveit, M.E. (eds.), The Commercial Storage of Fruits, Vegetables,
and Florist and Nursery. USDA Handbook 66. U.S. Department of
Agriculture, Agricultural Research Service, Beltsville, MD. http://www. (accessed October 15, 2013).
Sevillano, L., Sanchez-Ballesta, M.T., Romojaro, F., and Flores, F.B. 2009.
Physiological, hormonal and molecular mechanisms regulating chill-
ing injury in horticultural species. Postharvest technologies applied
to result its impact. Journal of the Science of Food and Agriculture 89,
Singh, K., Weaver, R.J., and Johnson, J.O. 1978. Effect of applications of
gibberellic acid on berry size, shatter, and texture of Thompson
­seedless  grapes. American Journal of Enology and Viticulture 29,
Sugar, D., and Basile, S.R. 2013. Integrated ethylene and temperature condi-
tioning for induction of ripening capacity in ‘Anjou’ and ‘Comice’ pears.
Postharvest Biology and Technology 83, 9–16.
Sun, Y., Chen, P., Duan, C., Tao, P., Wang, Y., Ji, K., Hu, Y., Li, Q., Dai, S.,
Wu, Y., Luo, H., Sun, L., and Leng, P. 2013. Transcriptional regulation of
genes encoding key enzymes of abscisic acid metabolism during melon
(Cucumis melo L.) fruit development and ripening. Journal of Plant
Growth Regulation 32, 233–244.
Tacken, E., Ireland, H., Gunaseelan, K., Karunairetnam, S., Wang, D., Shultz,
O., Bowen, J., Atkinson, R.G., Johnston, J.W., Putterill, J., Hellens, R.P.,
and Schaffer, R.J. 2010. The role of ethylene and cold temperature in
the regulation of the apple POLYGALACTURONASE1 gene and fruit
softening. Plant Physiology 153, 294–305.


Thompson, J., Kader, A., and Sylva, K. 1996. Compatibility chart for fruits and
vegetables in short-term transport or storage. Agriculture and Natural
Resources Publication 21560. University of California, Oakland.
Timbie, M., and Haard, N.F. 1977. Involvement of ethylene in the hardcore
syndrome of sweet potato roots. Journal of Food Science 42, 491–493.
Toivonen, P.M.A. 2003. Effects of storage conditions and postharvest pro-
cedures on oxidative stress in fruits and vegetables. In Hodges, D.M.
(ed.), Postharvest Oxidative Stress in Horticultural Crops. Food Products
Press, New York, pp. 69–90.
Toivonen, P.M.A. 2004. Postharvest storage procedures and oxidative stress.
HortScience 39, 938–942.
Toivonen, P.M.A. 2010. Postharvest physiology of vegetables. In Hui, Y.H.,
Sinha, N., Ahmed, J., Evranuz, E.Ö., and Siddiq, M. (eds.), Handbook of
Vegetables and Vegetable Processing. Wiley-Blackwell Publishing, Ames,
IA, pp. 199–220.
Toivonen, P.M.A., and DeEll, J.R. 2002. Physiology of fresh-cut fruits and veg-
etables. In Lamikanra, O. (ed.), Fresh-Cut Fruits and Vegetables: Science,
Technology, and Market. CRC Press, Boca Raton, FL, pp. 91–123.
Toivonen, P.M.A., and Hodges, D.M. 2011. Abiotic stress in harvested fruits
and vegetables. In Venkteswarlu, B., and Shanker, A.K. (eds.), Abiotic
Stress in Plants—Mechanisms and Adaptations. Intech Open Access
Publishers, Rijeka, Croatia, pp. 39–59.
Toivonen, P.M.A., Hampson, C.R., Stan, S., McKenzie, D.-L., and Hocking,
R. 2007. Factors affecting severity of bruises and degree of apparent
bruise recovery in a yellow-skinned apple. Postharvest Biology and
Technology 45, 276–280.
Vinocur, B., and Altman, A. 2005. Recent advances in engineering plant toler-
ance to abiotic stress: Achievements and limitations. Current Opinion in
Biotechnology 16, 123–132.
Wang, C.Y. 1982. Physiological and biochemical responses of plants to chill-
ing stress. HortScience 17, 173–186.
Wang, C.Y. 1993. Approaches to reduce chilling injury of fruits and vegeta-
bles. Horticultural Reviews 15, 63–95.
Wang, C.Y. 2004. Chilling and freezing injury. In Gross, K.C., Wang, C.Y.,
and Saltveit, M.E. (eds.), The Commercial Storage of Fruits, Vegetables,
and Florist and Nursery. USDA Handbook 66. U.S. Department of
Agriculture, Agricultural Research Service, Beltsville, MD. http://www. (accessed October 15, 2013).
Wang, C.Y., and Qi, L. 1997. Modified atmosphere packaging alleviates chill-
ing injury in cucumbers. Postharvest Biology and Technology 10, 195–200.
Wang, H., Qian, S., Ma, S., Zhou, Y., Patrick, J.W., Duan, X., Jiang, Y., and Qu,
H. 2013. Energy status of ripening and postharvest senescent fruit of
litchi (Litchi chinensis Sonn.). BMC Plant Biology 13, 55.
Wang, K.L.-C., Li, H., and Ecker, J.R. 2002. Ethylene biosynthesis and signal-
ing networks. The Plant Cell (Suppl.), S131–S151.

P o s t h a r v e s t P hy s i o l o g y o f F r u i t s a n d V e g e t a b l e s

Watada, A.E., Herner, R.C., Kader, A.A., Romani, R.J., and Staby, G.L. 1984.
Terminology for the description of developmental stages of horticul-
tural crops. HortScience 119, 20–21.
Watkins, C.B. 2008. Dynamic controlled atmosphere storage: A new technol-
ogy for the New York storage industry? New York Fruit Quarterly 16,
Watkins, C.B., Kupferman, E., and Rosenberger, D.A. 2004. Apples. In
Gross, K.C., Wang, C.Y., and Saltveit, M.E. (eds.), The Commercial
Storage of  Fruits, Vegetables, and Florist and Nursery. USDA Handbook
66. U.S.  Department  of Agriculture, Agricultural Research Service,
Beltsville, MD. (accessed
October 15, 2013).
Whitaker, B.D. 2004. Oxidative stress and superficial scald of apple fruit.
HortScience 39, 933–937.
Wills, R.B.H., Ku, V.V.V., Shohet, D., and Kim, G.H. 1999. Importance of low
ethylene levels to delay senescence of non-climacteric fruit and vegeta-
bles. Australian Journal of Experimental Agriculture 39, 221–224.
Wills, R.B.H., Warton, M.A., and Ku, V.V.V. 2000. Ethylene levels associ-
ated with fruit and vegetables during marketing. Australian Journal of
Experimental Agriculture 40, 465–470.
Zhang, C., Tanabe, K., Tani, H., Nakajima, H., Mori, M., and Sakuno, E. 2007.
Biologically active gibberellins and abscisic acid in fruit of two later-
maturing Japanese pear cultivars with contrasting fruit size. Journal of
the American Society for Horticultural Science 132, 452–458.
Zhang, Z., Fu, R., Huber, D.J., Rao, J., Chang, X., Hu, M., Zhang, Y., and Jiang,
N. 2012. Expression of expansin gene (CDK-Exp3) and its modulation
by exogenous gibberellic acid during ripening and softening of persim-
mon fruit. HortScience 47, 378–381.
Zubo, Y.O., Yamburenko, M.V., Selivankina, S.Y., Shakirova, F.M., Avalbaev,
A.M., Kudryakova, N.V., Zubkova, N.K., Liere, K., Kulaeva, O.N.,
Kusnetsov, V.V., and Börner, T. 2008. Cytokinin stimulates chloroplast
transcription in detached barley leaves. Plant Physiology 148, 1082–1093.
Zuo, J., Zhu, B., Fu, D., Zhu, Y., Ma, Y., Chi, L., Ju, Z., Wang, Y., Zhai, B., and
Luo, Y. 2012. Sculpting the maturation, softening and ethylene pathway:
The influences of microRNAs on tomato fruits. BMC Genomics 13, 7.

Chapter 3

Postharvest Quality
of Ornamental Plants
Fernando L. Finger, Tania P. Silva, Fernanda
F. Araujo, and Jose G. Barbosa
Federal University of Viçosa, Viçosa, Minas Gerais, Brazil

Abstract 82
3.1 Introduction 82
3.2 Quality Attributes in Ornamental Plants 83
3.3 Influence of Water Relations on Ornamental Longevity 85
3.4 Action of Ethylene on Ornamental Plant Quality 88
3.4.1 Inhibition of Ethylene Synthesis 91
3.4.2 Inhibition of Ethylene Action 93
3.4.3 Ethylene Absorbers 95
3.5 Role of Abscisic Acid, Gibberellins, and Cytokinins 96
3.6 Role of Calcium on Flower Senescence 97
3.7 Respiration 98
3.8 Temperature 99
3.9 Handling of Cut Flowers 102
3.10 Potted Plants 103
3.11 Conclusions and Future Perspectives 104
Acknowledgment 105
References 105


In this chapter, the behavior of cut flowers and potted ornamental plants is
studied regarding the physiological and environmental factors that affect
the rate of senescence and longevity. Water uptake by cut flowers, carbohy-
drate supply, and response of flowers to ethylene interact alone or together
to affect the length of the vase life. Depending on whether ethylene is the
main cause of flower senescence, particular handling is required to extend
vase life. Periodically, recuts of the flower stem or pulsing with sucrose
improves the water uptake and floret opening of the bird-of-paradise. Any
flower species or potted plants with high sensitivity to ethylene have a
much better shelf life when treated with inhibitors of ethylene action such
as 1-methylcyclopropene (1-MCP) or silver thiosulfate (STS). The orchid
Epidendrum ibaguense had a positive response by reducing flower abscis-
sion when treated with the aminoethoxyvivylglycine (AVG) inhibitor of
­ethylene synthesis. But in those flowers insensitive to ethylene presence,
water status and carbohydrate supply play a very important role in the
length of the vase life. Thus, pulsing solutions containing sugar or STS
may be effective in postponing earlier senescence in many flower species.
Based on the rate of leaf yellowing and abscission of fruits and leaves, treat-
ment of potted ornamental peppers with 1-MCP prolongs postproduction
life in indoor conditions. However, ethylene partially reversed the inhibition
of 1-MCP on leaf yellowing and abscission.

3.1 Introduction
Ornamental plants, in general, have limited vase or postproduction life;
therefore, the use of adequate postharvest practices of handling is essential
to maintain the quality and prolong the usefulness of the potted plant or
flower. The applied techniques should allow for transportation, and for the
majority of ornamental species, relatively short-term storage—at wholesale
or retail stores—is required before reaching the final consumer.
From the consumer’s point of view, quality is associated with the length
of the shelf life, either at home or in display areas. But the shelf life varies
from ornamental plant to plant, as well as the initial quality of the product
and previous treatments applied to extend the postharvest life. Particularly
in underdeveloped and developing countries, the production areas of orna-
mental plants are not far from the selling commercial centers, and even with-
out modern postharvest handling, the final consumer still receives products
of reasonable quality. However, with the rapid increase in urban population,
the production areas have moved farther away. Under such a scenario, the
use of new techniques for preservation is required, which must guarantee

Postharvest Quality of Ornamental Pl ants

quality to the florist and be satisfactory to the ­consumer. Among the s­everal
techniques available for ornamental plant storage, temperature, relative
humidity, and composition of the atmosphere must be addressed. In addi-
tion, florists have to incorporate treatments to reduce the water and tem-
perature stresses, avoid the deleterious effects of ethylene, and minimize
the influence of low-level radiation in the storage and display areas.
After harvesting, the cut flowers or the potted plants at the postpro-
duction phase are subjected to a series of abiotic stresses, including water
stresses, intense transpiration, exposure to ethylene, and development of
physiological disorders. Usually, leafy ornamental plants are more resis-
tant to senescence than flowers. Flowers, on the other hand, due to their
ephemeral nature and the reduced supply of organic substances, find that
the catabolic metabolism accelerates quickly, thus altering important phys-
iological processes, such as reduction of the water uptake rate, depletion
of respiratory substrates, and an increase of ethylene production and sen-
sitivity. Despite the important influence of ethylene in reducing the vase
life of many ornamental crops, ornamental plants may respond to other
hormones, such as abscisic acid, gibberellins, and cytokinins, to extend of
their postharvest life.
Like any fresh product, ornamental plants, once harvested or trans-
ferred to the display areas, maintain intense respiratory and transpiratory
activities. Respiration will deplete the reserves of organic substrates, which
are already limited, whereas intense transpiration will result in the fast wilt-
ing of leaves and flowers.
Production of carbon dioxide by the ornamental plants can be dimin-
ished by reducing the storage temperature, usually as low as possible, but
above the freezing point of the cells. Many ornamental plants, however,
originate from tropical and subtropical climates and are susceptible to
chilling injury, which restricts the length of shipping and storage at low
In this chapter, we address the different aspects involved in the pres-
ervation of potted ornamental plants and cut flowers, with special attention
to postharvest treatments available to diminish the rate of leaf and flower

3.2  Quality Attributes in Ornamental Plants

For most ornamental plants, there is no government-required quality
­g rading system. However, most florist associations voluntarily establish
grading standards, mainly for cut flowers and foliage. As a general rule, for
cut flowers, emphasis is given to the uniformity of the plant material, focus-
ing on the stem length and diameter and the presence of curved portions.


Regarding the flower itself, the most important attributes are the size of the
bud, stage of development, and lack of wilted or abscised petals.
Two important criteria must be achieved at harvest for any ornamen-
tal plant: a presence quality compatible with the consumer’s wishes and an
extended shelf life after harvest or postproduction of potted plants, which
will allow enough time for transportation, eventual storage, and final dis-
play. In order to reach these goals, ornamental plants have to be harvested
at a specific stage of development, as presented in Table 3.1.
The establishment of the developmental stage for harvesting a
­particular flower is based on the ability of its bud flower to open after har-
vest, its response to abiotic stresses, and the length of its shelf life.
The standard quality for potted plants depends on the plant s­ pecies.
For instance, for chrysanthemum and ornamental peppers, the plant canopy

Table 3.1  Stage of Flower Development at Harvest

Scientific Name Common Name Development at Harvest
Anthurium cultorum Anthurium Spadix 3/4 fully developed
Antirrhinum majus Snapdragon Inflorescence with 1/3 to 2/3 of
open florets
Cattleya hybrids Orchid Fully open flower
Delphinium ajacis Delphinium Inflorescence with 1/2 of open
Dendrobium spp. Dendrobium Almost fully open flowers
Dianthus caryophyllus Carnation From half to fully open flower
Eustoma grandiflorum Lisianthus Oldest flower from the stalk fully
Gerbera hybrids Gerbera When two outer disc rows
are open
Gladiolus hybrids Gladiolus When few florets are showing
Gypsophila Baby’s breath Flowers fully open
Heliconia spp. Heliconia Harvest at stage of final display
Iris hybrids Dutch iris Flower buds showing color
Lilium spp. Asiatic lily Lower buds showing color
Rosa hybrids Rose Beginning of the bud petal
to unfold
Strelitzia reginae Bird-of-paradise From first floret showing color
to fully open
Zinnia elegans Zinnia Almost fully open flower

Postharvest Quality of Ornamental Pl ants

should cover the whole surface area of the pot. Also, shorter plants with
a compact canopy have a better appearance than taller ones. But in pots
containing orchids and African violets, the presence of flowers in different
stages of opening and the absence of senescent flowers or leaves are more
important than the shape of the plant canopy.

3.3  Influence of Water Relations

on Ornamental Longevity
Harvested ornamentals plants, in particular cut flowers, usually present an
imbalance between water uptake and transpiration throughout the vase life,
which results in the wilting of petals and premature senescence. Several
causes can be attributed to the reduction of water absorption by the flow-
ers, including obstruction of the xylem by microorganisms, deposition of
pectin and phenols, and air embolism (Van Doorn, 1997). Cut carnations
and roses present a reduction of longevity due to an imbalance of water
status, which is related to the diminished capacity of water uptake by the
cut flower ­during the vase life. The reduction of the water uptake rate
accounts for the continuous drop of the fresh weight of the flower during
the vase life. Van Doorn et al. (1995) found the presence of Pseudomonas
spp., Acinetobacter calcoaceticus, and Alcaligenes spp. bacteria in a vase of
carnations after 10 days at room temperature. The presence of bacteria in
the water caused obstruction of water uptake by the stem and reduced the
longevity, which was inversely proportional to an increased bacterial count
in the vase water. However, regardless of the cause of vase blockage, the
hydraulic conductance is diminished throughout the vase life of the flower,
establishing disequilibrium between the water uptake and transpiration. In
Zinia elegans, Carneiro et al. (2002) observed that one of the first symptoms
of reduced water uptake by the stem was a decrease of its fresh weight,
which began within the first 24 h of harvest. However, when the base of
the stem was recut at a frequency of every 12 h, the weight loss rate was
diminished compared to that of the uncut flower control. This simple action
of recutting the base of the stem improved the flower water balance, keep-
ing the whole flower with a higher water content (Figure 3.1). But in flowers
like Eustoma grandiflorum, placed in similar environmental conditions, a
noticeable decrease was seen in the fresh weight of the whole stem over a
much longer period of time, after only 6 days in the vase (Liao et al., 2001).
Due to the rapid weight loss, Z. elegans flowers rapidly show symptoms of
wilting, in both petals and leaves; a delay of weight loss with concomitant
extension of its longevity was achieved by recutting the base of the stem. In
E. ­grandiflorum and cut roses, an extension of the longevity can be obtained
by adding 250 mg L −1 aluminum sulfate, a well-known antimicrobial agent.



Fresh weight (%)




0 20 40 60 80 100 120
Hours after harvest

Figure 3.1  Behavior of fresh weight in Zinnia elegans cut flowers recut ( )
and uncut ( ) at base of the stem every 12 hours. (From Carneiro, T.F. et al.,
Pesquisa Agropecuaria Brasileira 37, 1065–1070, 2002.)

Also, the inclusion of 200 mg L −1 8-hydroxyquinoline citrate or sulfate in

the vase solution extends the vase life of roses, due to less bacterial growth
in the water.
Once the bird-of-paradise is harvested, the content of water in the
flower keeps diminishing during the vase life, due to physical blockage at
the base of the stem. Apparently, the blockage of the xylem is due to the
high activity of peroxidase, a group of isozymes responsible for deposit-
ing complexes of oxidized phenolic compounds outside of the cell wall. In
this flower, the water status in the tissues can be maintained unchanged
by recutting the stem base by 2 cm every 2 days (Figure 3.2). Such a pro-
cedure keeps the relative water content of the sepal close to 93.9% during
the vase life of the flower, while in uncut stems the water content drops to
approximately 86% (Figure 3.2). As a result, the cutting of the stem base
extends the vase life of the flower by at least 1 day compared to that of
uncut ones. In other cut flowers such as chrysanthemum, xylem occlusion
in the base of the stem occurs due to the joint activity of peroxidase and
polyphenoloxidase; both enzymes are involved in the biosynthesis of lignin

Postharvest Quality of Ornamental Pl ants


Relative water contet (%)




0 2 4 6 8 10

Figure 3.2  Relative water content in the sepals of bird-of-paradise flower

recut every 2 days ( ) and uncut ( ) maintained at 25°C and 60% rela-
tive humidity. (From Campanha, M.M. et al., Revista Brasileira Horticultura
Ornamental 3, 27–31, 1997.)

and suberin (Van Doorn and Vaslier, 2002). But when inhibitors for either
enzyme were applied to vase water, a delayed blockage of the xylem vessels
was observed, resulting in a longer time until leaf wilting for the cut chry-
santhemum flower.
In roses, the vascular occlusion in the xylem is primarily associated
with the presence of bacteria in the cut at the base of the stem and by the
growing of bacteria in the water of the vase, blocking the water flow to the
flower due to the presence of extracellular polysaccharides and other degra-
dation products that originated from the dead bacteria (Van Doorn, 1997). In
such condition, the roses will develop several symptoms of water deficiency,
including lack of petal opening, bending of the stem neck below the flower,
and leaf wilting (Bleeksma and Van Doorn, 2003). But when roses were main-
tained in the vase in a solution containing 200 mg L −1 8-­hydroxyquinoline
sulfate alone or mixed with 30 g L −1 sucrose, the hydraulic conductance of
the stem was kept close to the initial level at harvest, and as consequence,


flower longevity was prolonged (Ichimura et al., 1999). Furthermore, the

addition of sucrose to the solution provided longer vase life by increasing the
carbohydrate content in the petals.
Hard water contains minerals that turn the water pH to alkaline,
which diminishes its movement through the plant tissues. Such a problem
can be solved by removing the minerals from the water with a deionizer or
by lowering the pH, making the water acidic. The pH of hard water should
be lowered to 3.5–4.0 by adding citric acid to the vase water.

3.4  Action of Ethylene on Ornamental Plant Quality

Cut flowers or potted plants are often exposed to several kinds of stresses
once they are harvested or moved from the field or greenhouses. Different
abiotic stresses may occur during shipping, storage in wholesale ware-
houses, retail stores, and in the final display areas, usually in indoor envi-
ronments. Exposing plants to ethylene may induce a variety of effects, from
inducing senescence and abscission of flowers and leaves to impacting the
rate of flower opening. Some of the effects of ethylene on ornamental plants
are shown in Table 3.2. The toxic symptoms of ethylene action vary consid-
erably according to the species, but always induce deleterious effects that
reduce the display life.

Table 3.2  Symptoms of Ethylene Action in Ornamental Cut and Potted Plants
Plant Symptoms
Alstroemeria Darkening, petal abscission
Asiatic lily Abscission of flower, inhibition of flower opening
Carnation Flower wilting
Chrysanthemum Accelerates flower, leaf senescence
Delphinium ajacis Flower abscission
Gypsophila Flower wilting
Iris Accelerates flower senescence
Orchids Flower abscission, senescence, epinasty, reddish petal
Ornamental Leaf yellowing; abscission of fruits, leaves, and flowers
Poinsettia Leaf and flower abscission, leaf epinasty
Roses Accelerates flower senescence, induces bud and petal
abscission, interruption of bud flower opening
Snapdragon Floret abscission

Postharvest Quality of Ornamental Pl ants

Ethylene is produced by any living plant tissue, but it is also a by-

product of modern industrial human activity. In most ornamental plants,
undesirable effects can be ethylene induced, including petal wilting and
abscission, epinasty, and chlorophyll loss in leaves. Woltering and Van
Doorn (1988) concluded that flower species could be classified in three
types as follows: wilting mediated by ethylene, wilting not mediated by eth-
ylene, and petal abscission mediated by ethylene.
Synthesis of ethylene in plants is initiated from the amino acid
­methionine as follows: L-methionine → S-adenosyl methionine (SAM) →
amino-1-cyclopropane-1-carboxylic acid (ACC) → ethylene. The formation
of ACC from SAM and the oxidation of ACC to ethylene are catalyzed by
ACC synthase (ACS) and ACC oxidase (ACO), respectively. These two
enzymes play a major role in the regulation of ethylene production, being
stimulated by abiotic and biotic stresses. According to the changes in eth-
ylene production, activity of ACS and ACO, and behavior of respiration
during the development of a flower, it can be identified as climacteric, non-
climacteric, and in some cases, with an intermediate behavior. Carnation,
orchids, and Delphinium flowers have a typically climacteric behavior, char-
acterized by the autocatalytic production of ethylene and a parallel increase
in CO2 production. In flowers like roses and Narcissus, a nonclimacteric
senescence occurs during the senescence. Contrary to climacteric flowers,
in a nonclimacteric flower, such as Grevillia cv. ‘Sylvia’, a poor correlation is
observed among ethylene production, respiration, and changes in the con-
tent of ACC during senescence (Setyadjit et al., 2004). Nevertheless, the
climacteric increase of ethylene production and respiration varies among
the different cultivars from the same flower species, as well as the response
and sensitivity to ethylene. During the flower senescence of carnation and
other climacteric flowers, the autocatalytic induction of ethylene produc-
tion observed in the petals is due to an increase in the expression of mRNAs
from the ACC synthase and ACC oxidase.
The senescence in the majority of climacteric flowers can be rapidly
induced by the pollination phenomenon, observed in carnation, orchids,
and Petunia. In these flowers, pollination increases ethylene production,
inducing petal wilting and abscission. Prevention of these deleterious ethyl-
ene-induced effects can be achieved by emasculation or by pretreating the
flowers with antiethylene action compounds. Pollination of the cut orchid
Phalaenopsis induced a rapid senescence, characterized by the wilting of
the petals (Porat et al., 1994b). These same authors found that in a control
not pollinated, the time for the incipient wilting of Phalaenopsis flowers at
room temperature was 19.2 days, similar to that of the emasculated flowers,
while in pollinated flowers, the time for the wilting dropped to 2.0 days.
The pollination of Phalaenopsis flowers induced senescence of the orchid
by increasing autocatalytic ethylene production, as well as enhancing the
sensitivity to ethylene.


Flower species have different degrees of sensitivity to ethylene,

v­ arying from insensitive, such as roses, bird-of-paradise, and Gladiolus, to
very sensitive, including carnation and orchids. Nevertheless, the degree of
sensitivity to ethylene in the same species varies from cultivar to cultivar,
as observed in roses.
As shown in Table 3.3, the number of important commercial cut
­flowers sensitive to the presence of ethylene is superior to the number of
flowers with low sensitivity or ethylene insensitivity.
Species of flowers considered to be highly sensitive to ethylene show
physiological responses in the presence of relatively low concentrations of the
hormone, usually between 0.1 and 1.0 µl L −1 air, when exposed for a period of
6–12 h. When a cut orchid Phalaenopsis is exposed to ethylene at a concentra-
tion of 0.1 µl L −1 air for 12 h, the longevity of the flower is reduced by 4.7%,
and it increases to 27.9% for a concentration of 0.5 µl L −1 (Porat et al., 1994b).
The action of ethylene in the senescence of the orchid Epidendrum
ibaguense is affected by the increase in ethylene concentration. A fast
increase in flower abscission and reduction of longevity was present when
the cut flowers were sprayed with different concentrations of ethephon
(Figure  3.3). At a concentration of 0.1 mg L −1 ethephon, an exponential
increase in the rate of abscission is observed, reaching saturation for the
response at concentrations of 100 mg L −1 ethephon or above. As a conse-
quence, the longevity dropped from 6.8 days for the unsprayed flowers to
3 days for the 100 and 1000 mg L −1 ethephon treatments (Figure 3.3).

Table 3.3  Degree of Sensitivity to Ethylene in Commercial Flower Species

High Low Insensitive
Alstroemeria Anthurium Strelitzia reginae
Consolida ajacis (Dephinium) Aspargus Roses
Iris Gerbera Gladiolus
Gypsophila paniculata Tulipa hybrida Sandersonia
Narcissus pseudonarcissus Waxflower Orchid (Cymbidium)
Orchids (Phalaenopsis, Chrysanthemum Buddleia davidii
Dendrobium, Epidendrum)
Petunia hybrida Cosmos bipinnatus
Carnation Cersis canadensis
Snapdragon Echinacea purpurea
Lathyrus odoratus
Hibiscus rosa-sinensis

Postharvest Quality of Ornamental Pl ants

25 8

Abscission (flower day −1)

Longevity (days)


0 200 400 600 800 1000
Ethephon (mg l−1)

Figure 3.3  Influence of ethephon on abscission ( ) and longevity ( ) of

orchid Epidendrum ibaguense. (From Moraes, P.J. et al., Acta Horticulturae
813, 565–570, 2009.)

As previously determined for other orchid flowers, Epidendrum

­ibaguense can be classified as a high ethylene-sensitive species. But not
all orchid species are sensitive to ethylene, such as Cymbidium (Table 3.3),
which shows no increase in flower abscission when it is exposed to
­ethylene. Nonetheless, flower discoloration is hastened in the presence of
the ­hormone (Van Doorn, 2002).

3.4.1  Inhibition of Ethylene Synthesis

The activity of ACC synthase can be reduced by applying solutions con-
taining rhizobitoxin analogues, such as aminoethoxyvivylglycine (AVG)
or aminooxyacetic acid (AOA). However, neither inhibitor can completely
knock down the production of ethylene. Such inhibitors have been tested
in both preharvest sprays and postharvest vase solutions and sprays, but
the efficiency of these treatments varies considerably among the flowers.
In orchid Dendrobium, the addition of 0.5 mM AOA to the vase solution
delayed the time to complete petal wilting and blocked the ethylene pro-
duction induced by the flower pollination (Porat et al., 1994a). On the other
hand, in Gypsophyla paniculata, another high ethylene-sensitive flower,


like the orchid Dendrobium, the use of commercial preservatives contain-

ing low concentrations of AVG had no effect compared to a control treat-
ment with no antiethylene substances (Newman et al., 1998). Commercial
AVG registered by the name of Retain® has been used in several countries
to control the natural senescence of ornamentals and fresh fruits and veg-
etables. When Retain was tested as a pulsing solution for at least 6 hours
or by spraying the inflorescence until it ran off cut Epidendrum ibaguense
flowers, a significant reduction of flower abscission was observed, which
resulted in extended vase life compared to that of the control treatment
(Table 3.4). Regardless of the way in which the AVG was applied, con-
centrations of 1.5 and 2.0 mM were more efficient in reducing the overall
flower abscission. Nonetheless, the spray was much more effective than
treating the flowers by pulsing. This difference in efficiency between the
two methods might be related to the faster uptake by the flower when a
spray solution is used.
The vase life of cut flowers, especially those sensitive to ethylene,
is extended by treating them with substances that can block the ethylene
action or reduce the hormone production. But in most cases, the inhibitors
of ethylene action seem to be more efficient in prolonging the longevity
of the flowers than the substances responsible for reducing the activity of
the key enzymes from ethylene synthesis, ACC synthase and ACC oxidase.
These obtain better results by the use of ethylene action blockers, in part
because the inhibitors of ACC synthase or ACC oxidase do not protect the
flower from action of exogenous ethylene produced from sources other than
the plant’s own tissue. Furthermore, as the AVG and AOA do not completely
knock down the production of ethylene, the remaining small ethylene pro-
duced by the flower can accelerate the senescence in sensitive ornamentals.
Maintaining roses in vase solution containing 5% sucrose plus 0.5,
1.0, 1.5, and 2.0 mM AOA significantly improved the longevity compared

Table 3.4  Percentage of Flower Abscission in Epidendrum ibaguense Pulsed

or Sprayed with AVG
AVG (mM) Pulsinga Spray
0.5 69.7 Ba 45.8 Ab
1.0 84.5 Aa 20.5 Bb
1.5 56.1 Ca 18.6 Cb
2.0 54.0 Da 18.5 Cb
Source: Mapeli, A.M. et al., Pesquisa Agropecuaria Brasileira 44, 258–262,
a Pulsing for at least 6 hours. Mean separation within columns by Tukey’s test;

values followed by the same uppercase and lowercase letters are not signifi-
cantly ­different at the 5% level.

Postharvest Quality of Ornamental Pl ants

to 5%  sucrose alone or control with distilled water, regardless of the

­concentration of AOA applied (Ketsa and Narkbua, 2001). However, the
authors did not measure the behavior of ethylene or CO2 production of
the  flowers in the presence of AOA. Based on their results, the addition
of AOA lowered the vase solution pH from 4.5 to 2.9, resulting in less growth
of bacteria in the solution, which might improve the water uptake and reduce
the development of bent neck.
Another inhibitor of ethylene production is acetylsalicylic acid (ASA),
which acts on the ACC oxidase enzyme. Fan et al. (1996) found that apple
ACC oxidase was strongly inhibited by ASA in both in vitro and in vivo
assays, as well as diminishing the fruit respiration. However, only a few
attempts have been made to evaluate the effectiveness of ASA in delay-
ing the deleterious effect of ethylene in cut flowers. When cut flowers of
Consolida ajacis were sprayed with 2 mM AOA or 20 mM ASA, they showed
no difference in vase life when compared to untreated flowers. However,
AOA and ASA had a slight effect in diminishing the initial rate of flower
abscission, which was not enough to extend the flower longevity (Finger
et al., 2004). In another work with roses, Brecheisen et al. (1995) found that
keeping the flowers in a solution containing 1% ASA significantly reduced
longevity when compared with tap water alone. In Consolida ajacis, neither
AOA nor ASA treatment was able to reduce the ethylene production of the
flowers, which might explain the lack of effect on the longevity.

3.4.2  Inhibition of Ethylene Action

The action of ethylene is efficiently inhibited by several inhibitors, including
silver ion, 1-methylcyclopropene (1-MCP), 2,5-norbornadiene, and nitrous
oxide. Among these inhibitors, 1-MCP seems to be the most promising
in blocking the ethylene, mainly due to its high affinity with the ethylene
receptor. Such ability makes the 1-MCP very effective at low concentra-
tions, ranging from 0.2 to 1 µl L −1. The commercial product is registered
by the name of EthylBloc ® for use in ornamental plants or SmartFresh® by
AgroFresh, Inc. 1-MCP is a worldwide commercialized growth plant regu-
lator, and it was classified as a nontoxic substance because similar natural
compounds are found in the atmosphere. 1-MCP is active at very low con-
centrations (Blankenship and Dole, 2003; PAN Pesticides Database, 2004).
Much work has been done in both ethylene-sensitive and -insensitive
flowers with 1-MCP. Usually the product is used just after harvest, fumigat-
ing the plants for 6–12 hours at room temperature, with different results
regarding its efficiency in blocking the ethylene action. The results vary
due to several endogenous and exogenous factors, including time between
harvest and the actual treatment, stage of development of the flower, con-
centration, and length of and temperature during the treatment.


Cut flowers of Consolida ajacis treated with 1-MCP had their longevity
extended significantly, even when the flowers were exposed to 100 mg L −1
ethephon after they had been exposed to 0.5 g m−3 SmartFresh for 6 h at room
temperature. When ethephon was applied 2 h before the use of 1-MCP, lower
longevity was observed than with 1-MCP alone or when 1-MCP was applied
after the flowers had been treated with ethephon. But for ethephon-treated
flower, a longer vase life was observed than for control fl
­ owers (Table 3.5).
Thus, the reduction in flower longevity when ethephon was applied before
1-MCP suggests that the latter did not occupy the entire available receptor
sites for ethylene.
In another work, Cameron and Reid (2001) found that 1-MCP had
a transitory effect to block the abscission of Pelargonium peltatum petals
induced by ethylene, and a second fumigation was needed to prevent flower
abscission. Thus, the reduction of flower longevity when ethephon was
applied before 1-MCP in Consolida ajacis and the transient effect observed
in P. peltatum flowers suggest that binding sites for ethylene were not com-
pletely occupied by 1-MCP, or new receptors sites were available by the
presence of exogenous ethylene. It is well known that the presence of ethyl-
ene is able to make the flowers more sensitive to its action.
In addition to 1-MCP, other similar cyclopropene compounds have
shown high effectiveness in blocking ethylene action. 1-Hexylcyclopropene
(1-HCP) and 1-octylcyclopropene (1-OCP) more effectively blocked the del-
eterious effects of ethylene in Kalanchoë blossfeldiana cv. ‘Alexandra’ flow-
ers than 1-MCP (Kebenei et al., 2003). The authors found that pretreatment
with 1-OCP was more effective than that with 1-MCP in delaying flower
senescence at a concentration of 200 nl L −1 for 6 hours, followed by exposi-
tion to 2 µl L −1 ethylene, while the 1-HCP pretreatment had an effect similar
to that of 1-MCP, but at a 5 or 10 times higher concentration.

Table 3.5  Effect of Ethephon and 1-MCP on the Longevity of Consolida

ajacis Flowers
Treatments Longevity (days)
Control 4.5 c
Ethephon 1.4 d
1-MCP 6.0 a
Ethephon + 1-MCP 5.2 b
1-MCP + Ethephon 6.0 a
Source: Santos, V.R. et al., Bragantia 64, 33–38, 2005.
Longevity: Days for 50% of flower abscission or wilting. Mean separation within
columns by Tukey’s test; values followed by a letter in common are ­similar at the
5% level.

Postharvest Quality of Ornamental Pl ants

Silver ion (Ag+) acts as an inhibitor of ethylene action. It is usually

sprayed or used in pulsing or in vase solutions to block the damaging
effects of ethylene, especially in flowers sensitive to ethylene. However, in
some flowers classified as having low or no sensitivity to ethylene, silver
ion may show beneficial effects when treated with silver thiosulfate (STS)
or AgNO3. In general, STS is more stable and has higher mobility within
the plant tissues than AgNO3; as a consequence, STS is considered less
phytotoxic and more effective at lower concentrations. STS is obtained by
mixing AgNO3 with sodium thiosulfate, with final concentrations varying
from 0.5 to 2.0 mM. To prepare a 0.1 M sodium thiosulfate stock solution,
mix 15.8 g of sodium thiosulfate into 500 ml of water. Separately, prepare
a 0.1 M silver nitrate stock solution by dissolving 16.9 g of silver nitrate
into 500 ml of water, keeping a 1:4 ratio of silver to thiosulfate. Pour the
silver nitrate solution onto the sodium thiosulfate under agitation. The STS
solution can be stored up to 1 month in the dark under refrigeration until
needed. Afterwards, the STS solution will lose its action due to silver pre-
cipitation and a new solution must be prepared.
Silver ion, mainly applied in the form of STS solution, has been used
commercially to prevent ethylene-induced senescence mainly in carnation.
But since silver is a heavy metal and considered a potential pollutant sub-
stance, other products have been tested to substitute Ag+ in the floriculture
industry. 1-MCP has been used as the most promising substitute of silver;
however, on some occasions STS is more efficient in extending the longev-
ity of ornamental crops than 1-MCP, in particular when the flowers are
stored in the absence of exogenous ethylene (Blankenship and Dole, 2003).
Pulsing roses with 1 mM AgNO3 or 1 mM STS for 3 hours delayed the petal
withering by 5–6 days compared to control flowers and extended the vase
life by 2.5 and 2.8 days, respectively (Son et al., 2003). Furthermore, due
to its action as a germicide at the base of the rose stem, silver ion was able
to maintain water potential at the flower peduncle throughout the vase life,
which resulted in improved water uptake by the stem. The treatment of
low ethylene-sensitive or ethylene-insensitive flowers does not always have
positive effects in extending their vase life. In tulips, for example, silver ion
showed no effect on the time, from the beginning of petal abscission, when
treated continuously with 0.5 mM STS (Sexton et al., 2000).

3.4.3  Ethylene Absorbers

Ethylene is an odorless and colorless gas that acts as a growth regulator
and harms horticultural crops. As an air pollutant, ethylene shows deleteri-
ous effects in ornamental plants in greenhouses, cold storage warehouses,
and inside display areas. Ethylene becomes active when the concentrations
in the atmosphere range from 0.1 to 1.0 µl L −1, and saturated responses


occur between the concentrations of 1 and 10 µl L −1. In order to lower the

presence of ethylene in the air, absorbers can be placed in the storage
room and in trucks during shipping. In addition to the ethylene originating
from the senescence of fruits, flowers, and leaves, ethylene exhausts come
from combustion engines, cigarette smoke, leaky gas lines, and biomass
The most common absorbers are alumina and potassium permanga-
nate, with the latter being largely used as an active ingredient in filters
and sachets. Potassium permanganate is able to oxidize ethylene, form-
ing carbon dioxide and water, and is active in cold storage warehouses,
refrigerated trucks, and at noncontrolled room temperatures. As an inac-
tive ingredient, it can be used in pieces of florist’s foam, dry silica gel, or
vermiculite. One can make a saturated solution of potassium permanganate
and mix it with the inactive ingredient. If necessary, allow it to dry in an
oven at 70°C; consider replacing the filter or sachet when the color changes
from purple to brown.

3.5  Role of Abscisic Acid, Gibberellins,

and Cytokinins
The use of abscisic acid, gibberellins, and cytokinins as flower preserva-
tives is still limited, because not too many flowers respond with positive
effects and because of the higher cost of the substances than of other com-
mercial compounds regularly applied to cut flowers.
Benzyladenine (BA) had showed different effects depending on the
species and cultivar of flower used. When tropical flowers were sprayed or
dipped in a solution containing 200 mg L −1 from 10 s to 1 min, it prompted
no improvement on the vase life of Strelitzia reginae and Zingiber spectabi-
lis (Paull and Chantrachit, 2001). Nonetheless, the BA improved the vase
life of Anthurium andraeanum, Heliconia psitacorum, Heliconia chartacea,
and Alpinea purpurata flowers from 1.5- to 2.5-fold. In a similar study,
Moraes et al. (2005) found that the response of Heliconia latispatha to spray
with BA was dependent on the concentration. The improvement on the
vase life increased linearly when the flowers were sprayed with 100, 200,
or 300 mg L −1 BA, and at highest concentration, the gain in longevity was
1.85-fold superior to that of the untreated flowers. Transgenic petunia flow-
ers overexpressing the isopentenyl transferase (ipt) gene from the cytoki-
nin biosynthesis had an improved longevity of 6 to 10 days longer than the
wild-type flowers (Chang et al., 2003). Also in this same study, the authors
found that the transformed plants were less responsive to exogenous eth-
ylene in inducing flower wilting, showing that the increase in cytokinin
­content caused by the overproduction of the ipt gene reduced the ­sensitivity
of the flowers to the ethylene.

Postharvest Quality of Ornamental Pl ants

Gibberellic acid (GA 3) was used in ornamental plants as an antagonist

to ethylene action, but its use as a preservative substance has shown other
effects in addition to reducing the flower senescence rate. Roses sprayed
with 1 mM GA 3 had suppressed the development of postharvest disease
caused by Botrytis cinerea by inhibiting the senescence-related changes in
the membrane permeability and protein degradation (Shaul et al., 1995).
The presence of exogenous ABA accelerated the leaf and flower
senescence of roses and carnation. When the daylily (Hemerocallis hybrid),
which is considered insensitive to ethylene, was treated with 100 µM ABA,
it caused premature accumulation of peroxidized lipids and induced the
activity of proteinase and RNAase, similar to the naturally occurring senes-
cence of the flower (Panavas et al., 1998). The authors suggest that ABA
plays an important role in the signal transduction events responsible for
the programmed death of daylily petal cells. In roses, Pompodakis et al.
(2004) found that the effect of ABA on flower longevity was dependent on
the pH used in the vase solution. In this work, the addition of 10 −5 M ABA at
pH 6 increased the vase life by inducing stomata closure in the presence or
absence of 1 mg L −1 AgNO3 as an antibacterial agent. But at alkaline pH 8, a
reduction of the vase life was observed, indicating that the beneficial effect
of ABA was eliminated by the microorganism growth present at high pH.

3.6  Role of Calcium on Flower Senescence

Pre- and postharvest applications of calcium have been applied to fresh
plant products to delay aging and postharvest decay and to control dis-
eases and physiological disorders. Flower and leaf senescence is delayed
by calcium treatment either before or after harvest, but the full mechanism
of its action still remains unclear. It is well known that calcium binds to
membrane phospholipids, protecting the membrane lipids from degrada-
tion, keeping the tissue function (Céour et al., 1992). Calcium also forms
Ca-pectates by cross-linking with COO− from the polygalacturonic acid at
medium lamella, increasing the rigidity of the plant cell wall. But the effec-
tiveness of calcium to improve the longevity of flowers such as cut roses
varies with the amount of calcium applied in the vase solution and also
by cultivar (Torre et al., 1999). In this work, the authors found that vase
solutions containing 1 mM CaCl 2 for the cv. ‘Baroness’ and 5 mM for the
cv. ‘Mercedes’ had delayed petal senescence, improved flower longevity,
enhanced opening of flower buds, postponed leakage of electrolytes, and
suppressed ethylene production during senescence. In addition, the CaCl 2
was able to keep the flower stems turgid for a longer period of time, retard-
ing the petal wilting when compared with untreated roses.
It has been observed that calcium also influences the development
of latent postharvest diseases in several fruits, vegetables, and flowers.


When cv. ‘Kiss’ roses were sprayed with 10 or 20 mM calcium sulfate mixed
with 0.01% Tween 20 until the solution ran off, applied 24 hours before har-
vest, postharvest infection with Botrytis cinerea was decreased during
display, thus increasing the vase life by at least 30% at room temperature
(Capdeville et al., 2005). In a similar work, Capdeville et al. (2003) found
that pulsing the same rose cultivar with 50 mM calcium sulfate for 15 hours
reduced the severity of gray mold caused by Botrytis cinerea by 88% and
increase the vase life.

3.7 Respiration
In general, flowers have high respiration rates compared to other horticul-
tural products, which might lead to carbohydrate depletion because flowers
are not adapted as a long-term storage organ. In addition, other deleterious
effects develop under high respiratory activity, including higher transpira-
tion rates and elevated ethylene production and action. The Q10 factor for
most of vegetables and fruits is close to 2, but in flowers, it might reach
values up to 8 (Wills et al., 1998). In cut Narcissus flowers, the respiration
rate in two commercial cultivars was exponentially increased when the
temperature was increased from 0°C to 12.5°C, given a Q10 close to 3.5.
The elevation of the temperature resulted in a negative linear relationship
between the respiratory activity and the vase life (Cevallos and Reid, 2000).
Similar respiration and vase life behavior was observed for cut flowers of
gerbera and sunflower when kept under temperatures ranging from 0°C to
20°C (Çelikel and Reid, 2002). Thus, for these flowers, at the temperature
conditions studied, the respiration can be used as an excellent index to pre-
dict the longevity of the vase life.
The respiration of flowers with high sensitivity to ethylene can be
reduced by applying inhibitors of its action, in particular STS, which acts
as a persistent inhibitor of ethylene action. Altman and Solomos (1995)
found that carnation flowers had no response to exogenous ethylene
regarding the development of petal wilting or an increase in respiration
when the vase solution contained 0.2 mM STS. The senescence of the
flowers in Consolida ajacis is associated with the increase in ethylene pro-
duction and respiration (Finger et al., 2004). When cut flowers of C. ajacis
were pulsed with 1 mM STS or 1 mM STS + 5% sucrose, the increase
of respiration was inhibited, thus prolonging the vase life of the flower
(Figure 3.4). However, by loading the flowers with sucrose alone, no influ-
ence on the longevity was detected, probably due to the weak effect of the
carbohydrate in lowering the respiration compared to the STS (Finger
et al., 2004). Similarly to sucrose pulsing, spraying the flowers with 2 mM
AOA or 20 mM ASA had no effect on the respiration or, as a consequence,
the flower longevity (Figure 3.4).

Postharvest Quality of Ornamental Pl ants

240 Control
5% sucrose
200 1 mM STS + 5% sucrose
Respiration (μl/g/h)




0 2 4 6 8 10 12 14 16
Days after harvest

Figure 3.4  Influence of pulsing with sucrose and STS and of spray with
AOA or ASA on respiration of Consolida ajacis flowers. (From Finger, F.L.
et al., Pesquisa Agropecuaria Brasileira 39, 533–537, 2004.)

3.8 Temperature
Temperature is the most important postharvest factor influencing the ­quality
and longevity of cut flowers and potted ornamental plants. Similar to any
fresh horticultural product, ornamental plants maintain high respiratory and
transpiratory activities to maintain their vital cell reactions. The intermedi-
ate organic molecules are used for the synthesis of new compounds and to
generate energy in the form of ATP. But respiration also produces heat as
a by-product, which increases when the temperature rises and accelerates
the aging process. The rate of senescence is dramatically reduced by cooling
the ornamental plants immediately after harvest. By far, the storage of cut
flowers at low temperatures has been used as the most adequate technique
for long-distance transport and, in many instances, for short-term storage
by the retail florists. The major effects of temperature on ornamental plant
storability and their posterior display are determined by the rate of respira-
tory substrate depletion and by the intensity of the transpiration rate and how
much production and action of ethylene will develop in the plants submitted
to temperature stress. Furthermore, all the previous effects of undesirable
storage temperatures will favor the development of postharvest diseases.


In potted roses, especially in those cultivars sensitive to ethylene,

the dropping of the bud flowers and yellowing of the leaves are extremely
reduced when the pots are transported or stored around 5°C and pretreated
with STS spray or 1-MCP fumigation. Storage at below room tempera-
ture and under low-irradiance inside light exposure, somewhere around
5–15  µmol m−2 s −1, induces rapid senescence in potted roses, the effects
of which can be minimized by lowering the temperature and blocking the
ethylene action. A similar influence of temperature and light intensity is
observed in chrysanthemum, petunia, and geranium potted plants.
Tropical and subtropical flowers are sensitive to chilling tempera-
tures; such species cannot be stored below a critical temperature ranging
from 0°C to 13°C. The development of chilling symptoms is dependent on
the species or variety sensitivity, stage of development, length of storage
under the chilling-inducing temperatures, and the actual temperature of
storage. For most horticultural products, temperatures near 5°C are usu-
ally more effective in developing chilling symptoms. Several commercially
important cut flowers are sensitive to chilling, such as bird-of-paradise,
orchids, heliconias, ginger, and Alpinea purpurata, among other ornamen-
tal species.
Several symptoms of chilling injury develop in ornamental plants,
making the product unsuitable for commercialization. One of the first symp-
toms to develop is an increase of metabolite leakage in the cells, indicating
the lack of semipermeability in the cell membranes. As a consequence, the
leakage of metabolites facilitates the growth of microorganisms, especially
of fungi-related diseases. Other visual symptoms and metabolic imbalance
reactions will develop when plants are under chilling injury–inducing tem-
peratures. Some of the most common symptoms include leaf and flower
wilt, intense water loss, water-soaked tissues, and leaf and flower discol-
orations. As a result of a metabolic imbalance reaction, there is acceler-
ated senescence and ethylene production and accumulation of ethanol and
The sensitivity and influence of storage length on the development
of chilling injury, and the subsequent influence on the vase life, were eval-
uated in bird-of-paradise flowers pulsed or not loaded with 40% sucrose
before storage at 10°C (Finger et al., 2003). Regardless of whether the
flowers were pulsed with sucrose before or after the cold storage, the
vase life at room temperature was reduced, with an increase in the length
of storage from 7 to 28 days (Table 3.6). In addition, the length of stor-
age reduced the total number of open florets at the end of the vase life
(Table  3.7). The most effective treatment in extending the flower lon-
gevity was achieved when the stems were pulsed after the cold storage
regardless of the number of weeks at 10°C, but when the flowers were
pulsed just before the cold storage, a fast decrease in poststorage vase

Postharvest Quality of Ornamental Pl ants

Table 3.6  Vase Life of Bird-of-Paradise Flowers after Pulsing with 40%
Sucrose for 24 Hours and 7, 14, 21, and 28 Days of Storage at 10°C
Vase Life (Days)
Treatments/Days of Storage
at 10°C 7 14 21 28
Distilled water 8.3 a 6.7 b 3.3 a 1.6 a
Pulsed before cold storage 5.6 b 4.2 c 2.0 b 0.7 b
Pulsed after cold storage 8.6 a 8.0 a 4.0 a 2.0 a
Source: Finger, F.L. et al., Acta Horticulturae 628, 863–867, 2003.
Note: Mean separation within columns by Tukey’s test; values followed by a
­letter in common are not significantly different at the 5.0% level.

Table 3.7  Number of Open Florets of Bird-of-Paradise Flowers after Pulsing

with 40% Sucrose for 24 Hours and 7, 14, 21, and 28 Days of Storage at 10°C
Number of Open Florets
Treatments/Days of Storage
at 10°C 7 14 21 28
Distilled water 1.7 b 1.6 b 1.2 a 0.5 b
Pulsed before cold storage 1.4 b 1.5 b 1.1 a 1.0 a
Pulsed after cold storage 2.5 a 2.0 a 1.2 a 0.7 ab
Source: Finger, F.L. et al., Acta Horticulturae 628, 863–867, 2003.
Note: Mean separation within columns by Tukey’s test; values followed by a
­letter in common are not significantly different at the 5.0% level.

life was observed (Table 3.6). Furthermore, by pulsing the flowers with
40% sucrose for 24 h after the 10°C storage, a higher number of open
florets was obtained, at least after 7 and 10 days of storage (Table 3.7).
The reduced vase life after 28 days of storage was associated with the
development of chilling symptoms, characterized by discoloration and
Penicillium spp. contamination in the flowers.
When the storage temperature for a particular ornamental species
is not known, determine the optimum temperature of storage by placing
the flowers at different temperatures, usually ranging from 0°C up to 15°C.
Most of the temperate ornamental crops are chilling insensitive and can be
stored between 0°C and 2°C. However, for plants that originate from tropi-
cal and subtropical regions, it is always important to consider their sensitiv-
ity to chilling injury. In the majority of cases, the symptoms of injury will
develop under the chilling-induced temperatures; if not, the plants have to
be moved to a higher temperature in order to show any disturbance.


3.9  Handling of Cut Flowers

In order to maintain the quality of cut flowers, it is necessary to address
the major factors that contribute to inducing or hastening the postharvest
deterioration during shipping, storage, and display. Understanding the sev-
eral factors that are directly involved in flower longevity will allow workers,
shippers, and wholesale and retail personnel to implement techniques to
maintain the quality of cut flowers:

1. Flower maturity: When flowers are harvested at their optimum

stage of development, a longer postharvest life will be achieved.
2. Stem cuts: Cut the stems with sharp and clean instruments and
immediately place them in buckets of water.
3. At the field: Keep the buckets with flowers in the shade and move
them to the packing house as soon as possible.
4. Temperature: After grading and cleaning the flowers, immediately
cool them to reduce the consumption of organic reserves, transpi-
ration and synthesis, and action of ethylene. If the flower is not
susceptible to chilling injury, cool the flowers to 2°C–4°C until
shipping or the product is moved to the display area.
5. Surface area: Flowers have a large surface area, which makes
them susceptible to intense water loss. To minimize wilting, store
the flowers in a cool chamber with relative humidity close to 95% or
cover them with perforated plastic bags.
6. Water: Use clean water at all times, and change it at least every
24 hours to avoid the growth of bacteria.
7. Buckets: Use light-colored buckets to make it easier to spot any
remaining dirt after cleaning.
8. Clean buckets: Wash the buckets with water containing 5% hypo-
chlorite before placing the flowers in them.
9. Hard water: Avoid the use of hard water with an alkaline pH in the
buckets. Use citric acid to lower the pH.
10. Tap water: Avoid the use of tap water due to the presence of fluo-
ride that can be toxic to some flowers, including roses and gladi-
olus. Instead of tap water, use rain water to keep the flowers.
11. Effects of ethylene: To reduce the undesirable effects of ethylene
on sensitive plants, the cold storage room should contain ethylene
absorber filters close to the evaporator fans. In cardboard boxes,
the use of ethylene absorber sachets can be effective in protecting
against ethylene action during shipping and storage.
12. Clean dying plant material: Clean up dying plant materials stored
in the cold storage chambers.
13. Moving methods: Avoid the use of gasoline- and propane-powered
carts around the storage areas.

Postharvest Quality of Ornamental Pl ants

14. Clear flame: Make sure that any burner close to the storage room
has a clear flame because a yellow-flame burn releases high
­ethylene-concentrated fumes.
15. Stability: Avoid excessive transit in the cold storage room due to
rapid changes in the temperature and relative humidity.

3.10  Potted Plants

The center of origin of the genus Capsicum is America, although it is cur-
rently grown in tropical and temperate regions of the planet (Casali and
Couto, 1984). The Capsicum peppers grown in pots have a dual use as an
ornamental plant and a source of edible fruits. The dual use of ornamental
plants as a source of beauty and as food adds value to the product, increas-
ing the financial return of the producer (Finger et al., 2012).
The proper choice of cultivar for the production of potted peppers is
vital for success in this type of venture. In general, producers of ornamen-
tal chili peppers use plants with smaller and more colorful fruits. Thus,
for the production of peppers in small pots, growers should seek cultivars
that result in low-growing plants when the fruits are ripe and colorful fruit
with intense and upright green foliage without symptoms of early senes-
cence and abscission of leaves and fruit. Two popular cultivars are the
‘Pyramid’ pepper (C. frutescens) and ‘Calypso’ (C. annuum). The variet-
ies used in pots can be of any species of the genus Capsicum, but with a
predominance of C. annuum, due to the great diversity of its form and the
color of its fruit.
There are several factors that affect the longevity of ornamental pep-
pers when transferred to indoor environments where there is a deficiency
of light and water, as well as an accumulation of ethylene. Preliminary stud-
ies have shown that ethylene triggers a series of negative reactions in pot-
ted peppers, among them the abscission of the fruit, leaves, and flowers as a
reaction to the growing sensitivity to gas. However, other effects are visible,
such as the accelerated degradation of chlorophyll and flower senescence.
The ethylene concentration required to cause these effects is dependent
on factors such as exposure time, temperature, stage of development, and
sensitivity of species or variety (Hoyer, 1996).
The ethylene synthesis can be stimulated by various environmen-
tal factors, such as temperature extremes, mechanical injuries and disor-
ders, the composition of the storage atmosphere, drought, and disease.
According to Khan (2006), the ethylene response may vary among species,
and each part of the plant has a different level of sensitivity to this hormone.
The most suitable inhibitors of ethylene action belong to the cycloolefin
class, which can be used for consumption. Among the cycloolefins, 1-MCP
has been widely used in the treatment of fruits, vegetables, and ornamental


Figure 3.5  Effect of ethylene and 1-MCP on the senescence of ornamental

pepper cultivar ‘Calypso’ (Capsicum annuum).

plants for control of fruit ripening and senescence of vegetables, cut ­flowers,
and ornamental potted plants (Blankenship and Dole, 2003).
Treatment with 1 g m−3 EthylBloc (0.14% of 1-MCP) for 6 h at a tem-
perature of 20°C to 25°C inhibited the action of ethylene on the abscission
of leaves of the ornamental pepper cultivar ‘Calypso’ (Segatto et al., 2013).
The application of 1-MCP extended the vase life of plants and completely
blocked the sites of ethylene action. Thus, once sprayed with 1-MCP, plants
do not exhibit abscission of leaves when exposed to ethylene (Figure 3.5).
There was no degradation of chlorophyll or leaf abscission in plants of the
cultivar ‘Calypso’, which were pretreated for 6 hours with 1 g m−3 EthylBloc
and subsequently exposed to an atmosphere containing 10 ml L −1 ethylene
at 48 h (Figure 3.5).

3.11  Conclusions and Future Perspectives

The longevity of flower and potted ornamental plants is determined by
the interaction of environmental, physiological, and genetic factors.
Water ­status, availability of carbohydrates, and ethylene action are deter-
minant factors in the development of senescence symptoms in most cut
flowers. In cut flowers and potted plants sensitive to ethylene, inhibitors

Postharvest Quality of Ornamental Pl ants

of ethylene action prolong vase and shelf life by reducing flower wilting
and abscission of leaves, flowers, and fruits. Furthermore, inhibitors of
ethylene synthesis seem to be less effective than inhibitors of its action
in delaying flower and leaf senescence. Additional data are required to
establish the critical levels of ethylene for the senescence of many orna-
mental plants. The role of abscisic acid, gibberellins, and cytokinins in
the senescence of flowers must be better understood before they can be
of any practical use.

FAPEMIG and CNPq provided scholarship and financial support.

Altman, S.A., and Solomos, T. 1995. Differential and morphological responses
of carnations pulsed or continuously treated with silver thiosulfate.
Postharvest Biology and Technology 5, 331–343.
Blankenship, S.M., and Dole, J.M. 2003. 1-Methylcyclopropene: A review.
Postharvest Biology and Technology 28, 1–25.
Bleeksma, H.C., and Van Doorn, W.G. 2003. Embolism in rose stems as
a result of vascular occlusion by bacteria. Postharvest Biology and
Technology 29, 334–340.
Brecheisen, S., Haas, H.P., and Röber, R. 1995. Influence of water quality and
chemical compounds on vase life of cut roses. Acta Horticulturae 405,
Cameron, A.C., and Reid, M.S. 2001. 1-MCP blocks ethylene-induced
petal abscission of Pelargonium peltatum but the effect is transient.
Postharvest Biology and Technology 22, 169–177.
Campanha, M.M., Finger, F.L., Cecon, P.R., and Barbosa, J.G. 1997. Water
relations of cut bird-of-paradise (Strelitzia reginae Ait.) inflorescences.
Revista Brasileira Horticultura Ornamental 3, 27–31.
Capdeville, G. de, Maffia, L.A., Finger, F.L., and Batista, U.G. 2003. Gray mold
severity and vase life of rose buds after pulsing with citric acid, sali-
cylic acid, calcium sulfate, sucrose and silver thiosulfate. Fitopatologia
Brasileira 28, 380–385.
Capdeville, G.D., Maffia, L.A., Finger, F.L., and Batista, U.G. 2005. Pre-harvest
calcium sulfate applications affect vase life and severity of gray mold in
cut roses. Scientia Horticulturae 103, 329–338.
Carneiro, T.F., Finger, F.L., Santos, V.R., Neves, L.L.M., and Barbosa, J.G.
2002. Influência da sacarose e da base da haste na longevidade de
inflorescências de Zinnia elegans. Pesquisa Agropecuaria Brasileira 37,


Casali, V.W.D., and Couto, F.A.A. 1984. Origem e botânica de Capsicum.

Informe Agropec 10, 8–10.
Çelikel, F.G., and Reid, M.S. 2002. Storage temperature affects the quality of
cut flowers from the Asteraceae. HortScience 37, 148–150.
Céour, F., Arul, J., Makhlouf, J., and Willemot, C. 1992. Delay of membrane
lipid degradation by calcium treatment during cabbage leaf senescence.
Plant Physiology 100, 1656–1660.
Cevallos, J.C., and Reid, M.S. 2000. Effects of temperature on the respiration
and vase life of Narcissus flowers. Acta Horticulturae 517, 335–341.
Chang, H., Jones, M.L., Banowetz, G.M., and Clark, D.G. 2003.
Overproduction of cytokinins in petunia flowers transformed with
PSAG12-IPT delays corolla senescence and decreases sensitivity to ethyl-
ene. Plant Physiology 132, 2174–2183.
Fan, X., Mattheis, J.P., and Fellman, J.K. 1996. Inhibition of apple
fruit  1-aminocyclopropane-1-carcoxylic acid oxidase and respiration
by ­acetylsalicylic acid. Journal of Plant Physiology 149, 469–471.
Finger, F.L., Carneiro, T.F., and Barbosa, J.G. 2004. Senescência pós-­colheita
de flores de esporinha (Consolida ajacis). Pesquisa Agropecuaria
Brasileira 39, 533–537.
Finger, F.L., Moraes, P.J., Barbosa, J.G., and Grossi, J.A.S. 2003. Vase life of
bird-of-paradise flowers influence by pulsing and term of cold storage.
Acta Horticulturae 628, 863–867.
Finger, L.F., Rêgo, E.R, Segatto, F.B., Nascimento, N.F.F., and Rêgo, M.M.
2012. Produção e potencial de mercado para pimenta ornamental.
Informatica Agropecuaria 33, 14–20.
Hoyer, L. 1996. Critical ethylene exposure for Capsicum annuum ‘Janne’ is
dependent on an interaction between concentration, duration and devel-
opmental stage. Journal of Horticultural Science 71, 621–628.
Ichimura, K., Kojima, K., and Goto, R. 1999. Effects of temperature,
8-hydroxyquinoline sulfate and sucrose on the vase life of cut rose flow-
ers. Postharvest Biology and Technology 15, 33–40.
Kebenei, Z., Sisler, E.C., Winkelmann, T., and Serek, M. 2003. Efficacy of
new inhibitors of ethylene perception in improvement of display life of
kalanchoë (Kalanchoë blossfeldiana Poelln.) flowers. Postharvest Biology
and Technology 30, 169–176.
Ketsa, S., and Narkbua, N. 2001. Effect of aminooxiacetic acid and sucrose on
vase life of cut roses. Acta Horticulturae 543, 227–234.
Khan, A.N. 2006. Ethylene Action in Plants. Springer, Amsterdam, Netherlands.
Liao, L.J., Lin, Y.H., Huang, K.L., and Chen, W.S. 2001. Vase life of Eustoma
grandiflorum as affected by aluminum sulfate. Botanical Bulletin of
Academia Sinica 42, 35–38.
Mapeli, A.M., Finger, F.L., Oliveira, L.S., and Barbosa, J.G. 2009. Longevidade
de inflorescências de Epidendrum ibaguense tratadas com aminoetoxi-
vinilglicina. Pesquisa Agropecuaria Brasileira 44, 258–262.

Postharvest Quality of Ornamental Pl ants

Moraes, P.J., Finger F.L., Mapeli A.M., Cecon P.R. and Barbosa J.G. 2009.
Growth and Flower Development of Epidendrum ibaguense Orchid.
Acta Horticulturae 813, 565–570.
Newman, J.P., Dodge, L.L., and Reid, M.S. 1998. Evaluation of ethylene
inhibitors for postharvest treatment of Gypsophila paniculata L.
HortTechnology 8, 58–63.
PAN Pesticides Database. 2004.
Panavas, T., Walker, E.L., and Rubinstein, B. 1998. Possible involvement of
abscisic acid in senescence of daylily petals. Journal of Experimental
Botany 49, 1987–1997.
Paull, R.E., and Chantrachit, T. 2001. Benzyladenine and the vase life of tropi-
cal ornamentals. Postharvest Biology and Technology 22, 303–310.
Pompodakis, N.E., Joyce, D.C., Terry, L.A., and Lydakis, D.E. 2004.
Effects of  vase solution pH and abscisic acid on the longevity of cut
‘Baccara’ roses. Journal of Horticultural Science and Biotechnology 79,
Porat, R., Borochov, A., and Halevy, A.H. 1994a. Pollination-induced changes
in ethylene production and sensitivity to ethylene in cut dendrobium
orchid flowers. Scientia Horticulturae 58, 215–221.
Porat, R., Borochov, A., Halevy, A.H., and O’Neil, S.D. 1994b. Pollination-
induced senescence of Phalaenopsis petals. Plant Growth Regulation 15,
Santos, V.R., Finger, F.L., Barbosa, J.G., and Barros, R.S. 2005. Influência do
etileno e do 1-MCP na senescência e longevidade de inflorescências de
esporinha. Bragantia 64, 33–38.
Segatto, F.B., Finger, F.L., Barbosa, J.G., Rêgo, E.R., and Pinto, C.M.F. 2013.
Effects of ethylene on the post-production of potted ornamental peppers
(Capsicum annuum L.). Acta Horticulturae 1000, 217–222.
Setyadjit, S., Joyce, D.C., Irving, D.E., and Simons, D.H. 2004. Development
and senescence of Grevillia ‘Sylvia’ inflorescences, flowers and flower
parts. Plant Growth Regulation 44, 133–146.
Sexton, R., Laird, G., and Van Doorn, W.G. 2000. Lack of ethylene involve-
ment in tulip tepal abscission. Physiologia Plantarum 108, 321–329.
Shaul, O., Elad, Y., and Zieslin, N. 1995. Suppression of Botrytis blight in
cut roses flowers with gibberellic acid: Effects of postharvest timing
of the gibberellin treatment, conidial inoculation and storage period.
Postharvest Biology and Technology 6, 331–339.
Son, K.C., Byoun, H.J., and Yoo, M.H. 2003. Effect of pulsing with AgNO3 or
STS on the absorption and distribution of silver and the vase life of cut
rose ‘Red Sandra’. Acta Horticulturae 624, 365–372.
Torre, S., Borochov, A., and Halevy, A.H. 1999. Calcium regulation of senes-
cence in rose petals. Physiologia Plantarum 107, 214–219.
Van Doorn, W.G. 1997. Water relations of cut flowers. Horticultural Reviews
18, 1–85.


Van Doorn, W.G. 2002. Effect of ethylene on flower abscission: A survey.

Annals of Botany 89, 689–693.
Van Doorn, W.G., and Vaslier, N. 2002. Wounding-induced xylem occlusion in
stems of cut chrysanthemum flowers: Roles of peroxidase and cathecol
oxidase. Postharvest Biology and Technology 26, 275–284.
Van Doorn, W.G., Witte, Y., and Harkema, H. 1995. Effects of high number of
exogenous bacteria on the water relations and longevity of cut carnation
flowers. Postharvest Biology and Technology 6, 111–119.
Wills, R., McGlasson, B., Graham, D., and Joyce, D. 2008. Postharvest: An
Introduction to the Physiology and Handling of Fruit, Vegetables and
Ornamentals. 4th ed. CABI, Wallingford, UK.
Woltering, E.J., and Van Doorn, W.G. 1988. Role of ethylene in senescence
of petals—morphological and taxonomical relationships. Journal of
Experimental Botany 208, 1605–1616.

Chapter 4

Physiology and
Molecular Biology of
Flower Senescence
Kenichi Shibuya and Kazuo Ichimura
National Agriculture and Food Research
Organization (NARO), Fujimoto, Tsukuba, Japan

Abstract 110
4.1 Introduction 110
4.2 Ethylene and Senescence of Cut Flowers 111
4.2.1 Ethylene Response of Cut Flowers 111
4.2.2 Types of Senescence in Cut Flowers with High
Ethylene Sensitivity 112
4.2.3 Ethylene in Petal-Wilting-Type Flowers 113
4.2.4 Ethylene in Petal-Abscission-Type Flowers 114
4.2.5 Ethylene Biosynthesis 114
4.2.6 Ethylene Signal Transduction 115
4.2.7 Acceleration of Flower Senescence by
Pollination 116
4.2.8 Acceleration of Flower Senescence by
Wounding 119
4.2.9 Effect of Temperature on Ethylene Production
and Perception 119


4.3 Plant Hormones Other than Ethylene in Flower

Senescence 120
4.3.1 Auxin 120
4.3.2 Gibberellin 120
4.3.3 Cytokinin 120
4.3.4 Abscisic Acid 121
4.3.5 Jasmonic Acid 122
4.4 Programmed Cell Death in Flower Senescence 122
4.4.1 Programmed Cell Death 122
4.4.2 Gene Expression during PCD in Flowers 123
4.4.3 Autophagy in Petal Senescence 124
4.5 Conclusions and Future Perspectives 124
References 125

Flower longevity is one of the most important traits of cut flowers. An under-
standing of the postharvest biology of flowers is needed to efficiently develop
postharvest technology. Ethylene is a key factor that regulates flower senes-
cence in many plant species. Ethylene biosynthetic and signaling pathways
have been well characterized, and it is now possible to improve the vase life
of some cut flowers by controlling the effects of ­ethylene. In contrast, regula-
tory mechanisms of senescence in flowers that show ­ethylene-independent
senescence remain largely unknown. Petal ­senescence is a highly regulated
developmental process and is considered to be a type of programmed cell
death, which is an actively regulated ­process. To identify key regulators of
programmed cell death during petal senescence, many studies have been
conducted. In this chapter, we outline studies on the postharvest physiology
and molecular biology of flowers, focusing on regulatory mechanisms for
petal senescence by ethylene and programmed cell death.

4.1 Introduction
Several postharvest techniques are applied to improve the vase life of many
cut flowers. These techniques have been developed based on knowledge
from basic studies of plant hormones, sugar, and water relations. Ethylene
plays a very important role in the senescence of many cut ­flowers, and this
plant hormone has been of primary interest in the postharvest biology of
flowers. Besides ethylene, other plant hormones play an important role in
flower senescence and the biochemical functions of biomembranes during
senescence. After the 1990s, with the advances in molecular b ­ iology, genes


involved in ethylene biosynthesis and ethylene signal transduction were

revealed in many ornamental plants. More recently, gene expression has
been analyzed comprehensively in several flowers, and numerous senes-
cence-related genes have been isolated. On the other hand, many reports
on the water relations of cut flowers have advanced the knowledge of post-
harvest physiology. Several reviews on the postharvest biology of flowers
have been published (Rogers, 2006, 2013; Tripathi and Tuteja, 2007; Van
Doorn and Woltering, 2008; Ichimura, 2010; Shibuya, 2012). In this chapter,
we review studies on the postharvest physiology and molecular biology of
flowers, focusing on regulatory mechanisms for petal senescence.

4.2  Ethylene and Senescence of Cut Flowers

4.2.1  Ethylene Response of Cut Flowers

Ethylene accelerates petal wilting or abscission of flower parts in some
plant species but has little effect in other plant species. Sensitivity to ethyl-
ene in cut flowers is generally evaluated based on whether signs of senes-
cence are observed after exposure to a constant concentration of ethylene.
Woltering and Van Doorn (1988) evaluated ethylene sensitivity in 96 plant
species from 24 families by treating with 3 ppm ethylene for 22–24 hours.
They reported that flowers of plant species belonging to the Asteraceae and
Iridaceae exhibit lower sensitivity to ethylene, whereas flowers belonging to
the Caryophyllaceae exhibit higher sensitivity, suggesting that differences
in ethylene sensitivity may depend on genetic factors. However, more recent
works found that longer exposure to ethylene results in accelerated flower
senescence in some species. For example, daffodil was classified as a flower
with very low ethylene sensitivity, but continuous treatment with ethylene
hastens flower senescence, and treatment with ethylene inhibitors delays
senescence (Hunter et al., 2004b). In some plant species, ethylene sensitiv-
ity of flowers increases with aging. Sepal abscission of Delphinium hybrid
cv. ‘Bellamosum’ flowers is not induced by ethylene treatment (40 µl L −1)
on the day of harvest, but it is induced if flowers are treated with ethylene
1 day after harvest (Ichimura et al., 2009a). Thus, time of treatment, con-
centration of ethylene, and age of flowers need to be considered in order to
evaluate the sensitivity to ethylene in cut flowers.
Sensitivity to ethylene varies greatly among plant species. In carna-
tion, 0.6 µl L −1 ethylene induces petal wilting within 12 hours (Wu et al.,
1991), while in rose it takes more than 2 days until flowers abscise in continu-
ous 1 µl L −1 ethylene (Ichimura et al., 2005). In chrysanthemum, little effect
is observed even when flowers are treated with 1 µl L −1 ethylene for more
than 10 days (Doi et al., 2003). Here, we have classified ethylene sensitivity
of cut flowers based on observation of their responses to ethylene (Table 4.1).


Table 4.1  Ethylene Sensitivity in Cut Flowers

Sensitivity Plant Species References
Very high Carnation (Nichols, 1968)
High Gypsophila paniculata (Woltering and Van Doorn, 1988)
Sweet pea (Shimizu-Yumoto and Ichimura, 2006a)
Delphinium (Ichimura et al., 2009a)
Dendrobium (Porat et al., 1994)
Vanda (Goh et al., 1985)
Relatively Campanula (Kato et al., 2002)
high Snapdragon (Ichimura et al., 2007)
Stock (Celikel and Reid, 2002)
Eustoma (Shimizu-Yumoto and Ichimura, 2009b)
Rose (Ichimura et al., 2005)
Blue star (Tweedia caerulea) (Hiraya et al., 2002)
Relatively Alstroemeria (Wagstaff et al., 2005)
low Daffodil (Hunter et al., 2004b)
Low Chrysanthemum (Doi et al., 2003)
Gladiolus (Serek et al., 1994)
Tulip (Sexton et al., 2000)
Lily (oriental hybrid) (Elgar et al., 1999)
Note:  Very high, petal wilting observed within 24 hours of treatment with ethylene
at less than 1 µl L−1; high, petal wilting or flower abscission observed within
24 hours of treatment with ethylene at 1–10 µl L−1; relatively high, petal wilt-
ing or flower abscission observed within 48 hours of treatment with ethylene
at 1–10 µl L−1; ­relatively low, acceleration of senescence symptoms eventually
observed at a significant level after ethylene or ethrel treatment; low, accelera-
tion of senescence symptoms not observed after ethylene treatment.

Some flowers, including Delphinium (Ichimura et al., 2009a), Eustoma

(Shimizu-Yumoto and Ichimura, 2009b), Portulaca (Ichimura and Suto,
1998), and Torenia (Goto et al., 1999), show an increase in ethylene sensitiv-
ity as flowers age. Intriguingly, ethylene sensitivity declines after harvest
in carnation flowers, which are highly sensitive to ethylene (Mayak and
Dilley, 1976b; Onozaki et al., 2004).

4.2.2  Types of Senescence in Cut Flowers

with High Ethylene Sensitivity
Senescence patterns of flowers that show high ethylene sensitivity can be
classified into two types: a petal-wilting type, in which ethylene induces
petal wilting, and a petal-abscission type, in which ethylene induces


abscission of petals and sepals. Abscission of flower parts is regulated by

­ethylene in many plant species, although there are exceptions, such as tulip
(Sexton  et al., 2000). Petal-wilting-type flowers include carnation, sweet
pea, and orchids, among others. Delphinium is a representative of a petal-
or sepal-abscission-type flower. In flowers such as sweet pea and blue star
(Tweedia caerulea), petals or whole flowers abscise after petals have wilted.
In these cases, senescence is generally classified by type based on whether
wilting or abscission occurs first.

4.2.3  Ethylene in Petal-Wilting-Type Flowers

Petal-wilting-type flowers can be further classified into two groups based
on patterns of ethylene production during flower senescence. Carnation
(Nichols, 1966), sweet pea (Mor et al., 1984), and Eustoma (Ichimura and
Goto, 2000) exhibit a clear increase in ethylene production during natural
(unpollinated) senescence. In contrast, ethylene synthesis does not rise
during natural senescence in some flowers, including Campanula (Kato
et al., 2002). In flowers in which ethylene synthesis rises during natural
senescence, treatment with ethylene inhibitors such as STS (anionic silver
thiosulfate complex) delays flower senescence. On the other hand, in flow-
ers in which ethylene synthesis does not rise during natural senescence,
such as Campanula (Kato et al., 2002), ethylene inhibitors have little effect
on the progression of senescence. In Campanula, ethylene production is
induced by pollination, resulting in hastened senescence. In this case, STS
treatment can suppress the increase in ethylene production and cancel the
effect of pollination on senescence (Kato et al., 2002).
Ethylene synthesis rises as flowers senesce, and treatment with ethyl-
ene inhibitors delays petal wilting in some petal-wilting-type flowers, indicat-
ing that ethylene regulates petal wilting in these flowers. Ethylene production
induced by ethylene itself is called autocatalytic ethylene production. The
climacteric-like rise in ethylene production during petal senescence in many
petal-wilting-type flowers is the result of autocatalytic ethylene ­production.
Since petal wilting is accompanied by autocatalytic ethylene production,
these two events are thought to be closely linked and inseparable. However,
in transgenic carnation, in which expression of an ACC oxidase gene is
­suppressed, exogenous ethylene treatment induces petal wilting but not auto-
catalytic ethylene production, suggesting that these two events may be under
the control of different pathways (Kosugi et al., 2000).
Ethylene is produced from virtually all organs of flowers, but its
­production varies among organs. Even within the same organ, production
differs among parts of the organ. In carnation petals, for example, the base
of the petals produces more than 10-fold more ethylene than the tip of the
petals on a fresh weight basis (Mor et al., 1985).


In carnation flowers, an increase in ethylene production starts in the

gynoecium prior to petals (ten Have and Woltering, 1997). Thus, gynoe-
cium-borne ethylene is thought to induce ethylene production in the pet-
als, leading to petal wilting. In fact, removal of the gynoecium has been
reported to suppress an increase in ethylene production in petals and to
dramatically delay petal senescence (Shibuya et al., 2000). However, the
role of the gynoecium in petal senescence needs to be clarified, since sev-
eral studies have reported different results (Mor et al., 1980; Sacalis and
Lee, 1987; Woodson and Brandt, 1991).
In Portulaca flowers, stamens produce more ethylene on a fresh
weight basis than petals or the gynoecium. When stamens are touched,
a large amount of ethylene is produced only from stamens, leading to
petal senescence (Shimizu and Ichimura, 2001). In contrast, wounding of
the gynoecium or petals does not accelerate petal senescence (Ichimura
and Suto, 1998). This suggests that in the case of Portulaca, the ethylene
­produced from stamens regulates petal senescence.

4.2.4  Ethylene in Petal-Abscission-Type Flowers

Petal-abscission-type flowers can be classified into two groups: flowers
that show autocatalytic ethylene production during senescence, such as
Delphinium (Ichimura et al., 2009a) and Torenia (Goto et al., 1999), and
flowers that do not show autocatalytic ethylene production during senes-
cence, such as geranium (Clark et al., 1997). In flowers of both types, an
increase in ethylene production is observed in tissues other than the petals.
In Delphinium flowers, ethylene is mainly produced from gynoe-
cia and receptacles, and its amount rapidly increases during senescence.
Wounding of gynoecia or receptacles induces ethylene production in these
organs and accelerates sepal abscission (Ichimura et al., 2009a). Since
sepals connect directly with receptacles in Delphinium, ethylene produced
from receptacles seems to play a major role in sepal abscission. In Digitalis
(Stead and Moore, 1979) and Torenia (Goto et al., 1999), ethylene produc-
tion rises in the gynoecium, while a little ethylene is produced in petals
during senescence. Thus, in these plants, ethylene that is produced from
the gynoecium is likely involved in petal abscission.

4.2.5  Ethylene Biosynthesis

The ethylene biosynthetic pathway in plants has been established, and
genes encoding key enzymes have been isolated (Yang and Hoffman, 1984;
Kende, 1993; Lin et al., 2009). Ethylene is synthesized through the follow-
ing pathway:


l -Methionine → S-adenosyl-l -methionine → 1-aminocyclopropane-

1-carboxylic acid (ACC) → ethylene

The last two reactions are catalyzed by ACC synthase and ACC
In vegetative tissues of plants, ACC synthase is considered to be a
rate-limiting step in ethylene biosynthesis. However, activities of not only
ACC synthase but also ACC oxidase rise during petal senescence of car-
nation, suggesting that both enzymes are responsible for the increase in
ethylene production (Woodson et al., 1992). In the gynoecium of carnation
flowers, on the other hand, ACC oxidase activity is constantly high during
senescence, and thus the increase in ethylene production in this tissue is
likely due to a rise in ACC synthase activity (Yangkhamman et al., 2007).
ACC synthase and ACC oxidase are encoded by multigene families,
and genes encoding these enzymes have been isolated in many ornamental
plant species. Three ACC synthase genes have been isolated from carna-
tion (Park et al., 1992; Henskens et al., 1994; Jones and Woodson, 1999a),
four from rose (Wang et al., 2004), three from snapdragon (Woltering et al.,
2005), two from geranium (Wang and Arteca, 1995), and two from petunia
(Lindstrom et al., 1999). ACC synthase genes are differentially regulated
in a tissue-specific manner. For example, among the three carnation ACC
synthase genes, DcACS1 is most abundant in petals, while DcACS2 and
DcACS3 are preferentially expressed in styles (Jones and Woodson, 1999a).
For ACC oxidase, two genes have been isolated from carnation
(Wang and Woodson, 1991; Norikoshi et al., 2008), one from rose (Xue
et al., 2008), one from snapdragon (Woltering et al., 2005), one from
Phalaenopsis (Nadeau et al., 1993), five from tulip (Momonoi et al., 2007),
four from ­petunia (Tang et al., 1993), and three from geranium (Clark
et al., 1997). ACC oxidase genes are regulated in a spatial- and temporal-­
specific m­ anner. In carnation, DcACO1 is clearly upregulated in petals dur-
ing ­senescence, while DcACO2 is constantly expressed in the gynoecium
(Norikoshi et al., unpublished data). Differential expression of ACC oxi-
dase genes has also been reported in petunia (Tang et al., 1994).

4.2.6  Ethylene Signal Transduction

Ethylene signaling is mediated by a complex multicomponent pathway
(Lin et al., 2009). ETR1 (ETHYLENE RESPONSE1), which encodes a His
kinase with homology to bacterial two-component regulators, has been
identified as an ethylene receptor (Chang et al., 1993). Five ethylene recep-
tor genes have been cloned from Arabidopsis thaliana (Hua et al., 1995;
Sakai et al., 1998). Analysis of loss-of-function mutations in multiple recep-
tors has shown that these receptors are negative regulators of ethylene


responses (Hua and Meyerowitz, 1998). The receptors act through CTR1
(CONSTITUTIVE TRIPLE RESPONSE1), which is homologous to the
Raf family of Ser/Thr kinases and negatively regulates ethylene signaling
(Kieber et al., 1993; Huang et al., 2003). Downstream of the receptor–CTR1
complex is EIN2 (ETHYLENE INSENSITIVE2), which has homology to
Nramp metal transporters and positively regulates signaling (Alonso et al.,
1999). EIN2 undergoes proteasome-mediated turnover (Qiao et al., 2009),
and ethylene stimulates phosphorylation-dependent cleavage and nuclear
movement of a carboxyl-terminal EIN2 fragment (Qiao et al., 2012). Toward
the end of the signaling pathway, ethylene responses are mediated by tran-
scription factors including EIN3 (ETHYLENE INSENSITIVE3) and ERF1
(ETHYLENE RESPONSE FACTOR1) (Chao et al., 1997; Solano et  al.,
1998). Regulation of the stability of EIN3 protein is a key aspect of the ethyl-
ene response (Guo and Ecker, 2003; Yanagisawa et al., 2003).
Genes encoding ethylene receptors have been isolated in several
ornamental plant species, for example, three in carnation (Shibuya et al.,
2002), five in rose (Müller et al., 2000a, 2000b), one in chrysanthemum
(Narumi et al., 2005), two in Delphinium (Kuroda et al., 2003; Tanase and
Ichimura, 2006), four in petunia (Wang and Kumar, 2007), two in geranium
(Dervinis et al., 2000), and two in gladiolus (Arora et al., 2006). Genes
have been reported that encode CTR1 in rose (Müller et al., 2002) and
Delphinium (Kuroda et al., 2004), EIN2 in petunia (Shibuya et al., 2004),
and EIN3 in carnation (Waki et al., 2001; Iordachescu and Verlinden, 2005),
rose (Müller et al., 2003), and petunia (Shibuya and Clark, 2006).
Ethylene receptors are negative regulators of ethylene signaling, and
depletion of the receptors is considered to lead to an increase in ethylene
sensitivity (Hua and Meyerowitz, 1998). However, it has been reported that
the transcript levels of ethylene receptors in rose are higher in cultivars with
higher ethylene sensitivity in rose (Müller et al., 2000a). The inconsistency of
the relationship between levels of transcript for ethylene receptors and ethyl-
ene sensitivity might be explained by posttranslational regulation of ethylene
receptor proteins. Kevany et al. (2007) showed that protein levels for ethylene
receptors in tomato are not correlated with corresponding transcript levels
during fruit development and that the receptors are rapidly degraded in the
presence of ethylene. Thus, the level of ethylene receptor proteins seems
to be responsible for ethylene sensitivity. Similar regulation of the ethylene
response is expected in flower senescence. Analysis of ethylene receptors at
the protein level is necessary in the study of flower senescence.

4.2.7  Acceleration of Flower Senescence by Pollination

Pollination accelerates petal wilting and abscission of flower parts in spe-
cies that show relatively high ethylene sensitivity, including orchids (Burg


and Dijkman, 1967; Porat et al., 1994), petunia (Gilissen and Hoekstra,
1984; Whitehead et al., 1984), Eustoma (Ichimura et al., 1998; Ichimura
and Goto, 2000), carnation (Larsen et al., 1995; Jones and Woodson, 1997),
Campanula (Kato et al., 2002), Delphinium (Okamoto and Ichimura, 2010),
and Torenia (Goto et al., 1999). In particular, flower longevity of orchid spe-
cies is dramatically shortened by pollination. In daffodil flowers, which
show relatively low ethylene sensitivity, pollination also accelerates flower
senescence (Hunter et al., 2004b).
Pollination raises ethylene production from flowers, and treatment
with inhibitors of ethylene biosynthesis or ethylene perception cancels
the acceleration of flower senescence in many plant species (Stead, 1992).
Furthermore, transgenic petunia plants with reduced ethylene sensitiv-
ity due to suppressed expression of PhEIN2 exhibit significant delays in
petal senescence after pollination (Figure 4.1) (Shibuya et al., 2004). These
observations suggest that ethylene is a primary signal for pollination-­
accelerated flower senescence.
In Eustoma (Shimizu-Yumoto and Ichimura, 2006b), petunia (Gilissen,
1977), and Digitalis (Stead and Moore, 1979), a greater amount of pollen
­pollinating the stigma results in faster progression of flower senescence.
In  this case, a greater concentration of ethylene inhibitors is needed to
­counter the effect of pollination in flower senescence.
Compatibility between pollen and gynoecium involves the accel-
eration of flower senescence by pollination. In self-incompatible petunia,
compatible pollination accelerates the second peak of ethylene synthesis,
leading to hastened petal senescence, while incompatible pollination does

Figure 4.1  Petal senescence in transgenic petunia with reduced ethylene

sensitivity. Flower of a wild-type plant 2 days after pollination (left) and
flower of transgenic plant with reduced PhEIN2 expression 8 days after pol-
lination (right). The petals of transgenic plants are still turgid even after the
ovaries have fully expanded. (From Shibuya, K. et al., Plant Physiology 136,
2900–2912, 2004.)


not accelerate the second peak of ethylene synthesis and does not lead to
hastened petal senescence (Singh et al., 1992). Similar phenomena have
been observed in carnation (Larsen et al., 1995).
In pollinated orchid flowers, ethylene sensitivity rises prior to an
increase in ethylene production. This rise in ethylene sensitivity is not
suppressed by an ethylene synthesis inhibitor, AOA, suggesting that the
increase in ethylene sensitivity is regulated independently from e­ thylene
synthesis (Porat et al., 1994, 1995a). In addition, treatment with STS, an eth-
ylene perception inhibitor, suppresses the increase in ethylene ­production.
These observations suggest that the dramatic rise in ­ethylene production
after pollination in orchid flowers may be caused by the perception of a rela-
tively low level of ethylene as a result of an increase in ethylene sensitivity.
Detailed analyses of ethylene production patterns after pollination
have been conducted in several flower species. Petunia flowers produce
ethylene in styles within 3 h and then in petals at 2–3 days after pollination
(Hoekstra and Weges, 1986; Singh et al., 1992). In carnation, an increase
in ethylene production is observed at around 12 hours in the stigma and at
24 hours in ovaries, and then at 36 hours in petals after pollination (Jones
and Woodson, 1999b). In Phalaenopsis, ethylene increases at 12 hours in the
stigma and column, at 24 hours in the labellum, and after 48 hours in the
perianth after pollination (O’neill et al., 1993).
It has been reported that in pollinated Phalaenopsis flowers, the accu-
mulation of mRNA for ACC synthase and ACC oxidase increases in the
gynoecium, while mRNA for ACC oxidase is not detectable in the perianth
(O’Neill et al., 1993). Based on this observation, the following model has
been proposed: in the perianth, ACC is translocated from other flower parts,
such as the gynoecium, and is converted to ethylene (O’Neill et al., 1993).
However, mRNA for ACC synthase has since been detected in the perianth
using real-time reverse transcription polymerase chain reaction (RT-PCR),
a more sensitive technique, and the level of PhalACS1 mRNA increases in
the perianth after pollination (Ichimura and Niki, unpublished data).
In petunia, when the style is removed within several hours of pol-
lination, petal senescence is not accelerated by pollination (Gilissen and
Hoekstra, 1984). This observation suggests the existence of mobile signals
that induce petal senescence after pollination. Candidates for the mobile
signals are ACC and ethylene. When radiolabeled ACC is applied to the
stigmas of carnation flowers, radioactive ethylene is produced both by the
gynoecia and by the petals, suggesting that ACC may be a signaling com-
pound transported from the stigmas to the petals (Reid et al., 1984). In con-
trast, radiolabeled ACC applied to the stigma is largely immobile in flowers
of Cymbidium (Woltering et al., 1995) and petunia (Woltering et al., 1997).
Furthermore, in petunia, eluates collected from the styles were able to
induce petal senescence, but ACC was not detected in the eluates (Gilissen
and Hoekstra, 1984). These observations suggest that ACC is not a mobile


signaling factor for petal senescence in these plant species. In addition to

chemical compounds, electrical signaling has been proposed to be involved
in the induction of petal senescence after pollination (Spanjers, 1981;
Fromm et al., 1995).

4.2.8  Acceleration of Flower Senescence by Wounding

In plant species in which flower senescence is hastened by pollination,
wounding of gynoecia often accelerates flower senescence. Wounding of
the stigma results in accelerated petal senescence in Eustoma (Ichimura
and Goto, 2000) and petunia (Lovell et al., 1987; Whitehead et al., 1984),
and hastened petal abscission in Torenia (Goto et al., 1999). In Delphinium,
wounding of ovaries or receptacles accelerates abscission of flowers
(Ichimura et al., 2009a).
In wounding-induced flower senescence, ethylene production is has-
tened, as observed in pollination-induced flower senescence. However,
the pattern of ethylene synthesis induced by wounding differs from that
induced by pollination in petunia (Whitehead et al., 1984; Hoekstra and
Weges, 1996). In pollinated petunia flowers, ethylene is exclusively pro-
duced by the stigma and style region, whereas wounding of the stigma
induces ethylene production both by the stigma and style region and by the
remaining flower parts (Woltering et al., 1997). These observations sug-
gest that the mechanisms that accelerate petal senescence are different in
pollination- and wound-induced petal senescence.

4.2.9  Effect of Temperature on Ethylene

Production and Perception
Ethylene production and perception are affected by temperature. In carna-
tion, the time to induction of petal senescence by exogenous ethylene is
prolonged under lower temperatures, suggesting that ethylene sensitivity
declines as temperature falls (Barden and Hanan, 1972). However, it is not
clear whether this decline in ethylene sensitivity is a unique phenomenon
among other physiological responses to low temperature, such as a decline
in respiration. In contrast, lily flowers, which have relatively low ethylene
sensitivity, show an increase in ethylene sensitivity when flowers are kept
at low temperature (Song and Peng, 2004).
Ethylene synthesis is suppressed by high temperature in fruits such
as tomato and apple (Lurie, 1998). In carnation flowers, high temperature
suppresses ethylene synthesis (Yangkhamman et al., 2005, 2007; Shibuya
and Ichimura, 2010). When cut flowers of carnation ‘Barbara’ are kept at


more than 32°C, ethylene production from the flowers is suppressed,

­resulting in a delay in petal wilting. However, flowers kept at such a high
temperature show inhibited petal growth and fading of color.

4.3  Plant Hormones Other than

Ethylene in Flower Senescence
Plant hormones include auxin, gibberellin, cytokinin, abscisic acid (ABA),
brassinolides, and jasmonic acid. Salicylic acid and strigolactones are also
considered plant hormones. Of these, auxin, gibberellin, cytokinin, abscisic
acid, and jasmonic acid have been analyzed in studies on flower senescence.

4.3.1 Auxin
Auxin at relatively high concentrations induces ethylene synthesis. In
carnation, treatment with indoleacetic acid or 2,4-dichlorophenoxyacetic
acid, a synthetic auxin, induces ethylene production, leading to hastened
petal senescence (Sacalis and Nichols, 1980; Wulster et al., 1982). Auxin
at high concentrations seems to induce ACC synthase activity, resulting in
an increase in ethylene production. In contrast, application of 1-naphtha­
leneacetic acid, another synthetic auxin, suppresses abscission of bougain-
villea flowers (Chang and Chen, 2001).

4.3.2 Gibberellin
Gibberellin treatment prolongs flower longevity by suppressing ethylene
synthesis in carnation (Saks et al., 1992). It also prolongs the vase life of
daffodil, which has relatively low ethylene sensitivity (Ichimura and Goto,
2002). Since it is difficult to accurately quantify the gibberellin content in
plants, little is known of the distribution and movement of gibberellins in
cut flowers.

4.3.3 Cytokinin
Cytokinin generally delays senescence in plants. In rose (Mayak and
Halevy, 1970) and carnation (van Staden et al., 1987), cytokinin-like activity
decreases in senescent flowers. Application of benzyladenine, a synthetic
cytokinin, to presenescent cut carnation flowers delays petal senescence
(Mayak and Dilley, 1976a; Eisinger, 1977; Cook et al., 1985).


Cytokinin treatment prevents the conversion of exogenously applied

ACC to ethylene in carnation petals, and ethylene treatment does not
induce endogenous ethylene synthesis in the petals of flowers treated with
­c ytokinin (Mor et al., 1983; Cook et al., 1985). However, cytokinin treatment
does not inhibit ethylene synthesis in flowers in which a rise in ­ethylene syn-
thesis has already started (Mor et al., 1983). These observations suggest
that cytokinin does not directly inhibit the activities of enzymes involved
in e­ thylene synthesis, but it may suppress the synthesis of enzymes or
decrease ethylene sensitivity.
A synthetic cytokinin, thidiazuron, has been reported to improve
the vase life of iris flowers, which have relatively low ethylene sensitivity
(Macnish et al., 2010). In transgenic petunia expressing bacterial ipt, which
encodes a key enzyme of cytokinin synthesis, petal senescence is delayed
as a result of an increase in cytokinin content (Chang et al., 2003). In these
transgenic plants, the peak of ethylene production from flowers is delayed,
ethylene sensitivity declines, and ABA content decreases.
Cytokinin is likely involved in the regulation of flower senescence
in many plant species, regardless of the degree of ethylene sensitivity.
However, it is largely unknown whether changes in cytokinin content occur
in senescing tissues.

4.3.4  Abscisic Acid

ABA is generally considered to be a plant hormone that accelerates
senescence. ABA content in petals increases as the petals senesce in
rose (Borochov et al., 1976b) and carnation (Eze et al., 1986; Hanley and
Bramlage, 1989). In carnation, ABA treatment accelerates ethylene produc-
tion and increases ethylene sensitivity (Mayak and Dilley, 1976a, 1976b).
On the other hand, ethylene treatment increases ABA content in carnation.
In flowers with low ethylene sensitivity, such as daylily (Panavas
et al., 1998) and daffodil (Hunter et al., 2004a), ABA treatment also hastens
flower senescence. In daffodil, ABA concentration in the perianth increases
as flowers senesce (Hunter et al., 2004a).
ABA treatment of cut roses with leaves removed accelerates petal
senescence, while it delays senescence when the leaves are not removed
(Halevy et al., 1974; Borochov et al., 1976a). This observation is explained
as follows: when the leaves are removed, ABA directly affects the petals
and hastens petal senescence; in cut roses with leaves, ABA suppresses
transpiration from the leaves, resulting in improved water uptake, leading
to the prolongation of vase life. In Eustoma, ABA treatment extends vase
life by improving the water conditions of cut flowers (Shimizu-Yumoto and
Ichimura, 2009a).


ABA generally accelerates flower senescence, but it sometimes

improves the vase life of cut flowers by suppressing transpiration, leading
to better water conditions.

4.3.5  Jasmonic Acid

There are few reports on the effect of jasmonic acid on flower senescence.
Treatment with jasmonic acid ethyl ester accelerates flower senescence in
petunia (Porat and Halevy, 1993), Phalaenopsis, and Dendrobium (Porat
et al., 1995b). Since inhibitors of jasmonic acid synthesis have been devel-
oped, these inhibitors can be used to delay flower senescence.

4.4  Programmed Cell Death in Flower Senescence

4.4.1  Programmed Cell Death

Flower longevity is species specific and is thought to be genetically con-
trolled. Treatment with cycloheximide, a protein synthesis inhibitor, delays
flower senescence in many plant species, including carnation (Ichimura
et al., 2009b), gladiolus (Jones et al., 1994; Yamane and Ogata, 1995), and
Japanese morning glory (Yamada et al., 2009). The result of cycloheximide
treatment in gladiolus is shown in Figure 4.2. In Japanese morning glory,
actinomycin D, which inhibits RNA synthesis, also delays petal senes-
cence (Yamada et al., 2009). Furthermore, suppression of expression of the

Figure 4.2  Effect of cycloheximide, a protein synthesis inhibitor, on flower

senescence of gladiolus flowers. Detached flowers were placed in distilled
water (left) or 1 mM cycloheximide for 4 days.


gene for deoxyhypusine hydroxylase, which is involved in the synthesis of

eukaryotic translation initiation factor 5A (eIF5A), results in delayed petal
senescence in carnation (Hopkins et al., 2007). Based on these observa-
tions, flower senescence is considered to be a type of programmed cell
death (PCD), which is an actively regulated process.
A typical hallmark of PCD in animal cells is the degradation of nuclei
and DNA. During petal senescence, the first sign of PCD is chromatin
condensation. In petal cells of petal-wilting-type flowers, including petunia
(Xu and Hanson, 2000; Langston et al., 2005) and Japanese morning glory
(Yamada et al., 2006), DNA degradation is observed during s­ enescence.
In  contrast, DNA degradation is not observed in petal cells of Prunus
yedoensis, a type of cherry tree, which is a petal-abscission-type flower, at
the point of petal abscission (Yamada et al., 2007b), suggesting that PCD
may not occur in this flower until the petals abscise.

4.4.2  Gene Expression during PCD in Flowers

Changes in gene expression during flower senescence have been analyzed
by microarrays or subtraction assays in several species, including Iris
(Van Doorn et al., 2003), Japanese morning glory (Yamada et al., 2007a),
Alstroemeria (Breeze et al., 2004), Mirabilis jalapa (Xu et al., 2007), carnation
(Hoeberichts et al., 2007), daffodil (Hunter et al., 2002), and daylily (Panavas
et al., 1999). In petunia, proteomic analysis has also been conducted and pro-
tein changes have been profiled during petal senescence (Bai et al., 2010).
Genes that are commonly upregulated during flower senescence in these
plant species are genes encoding enzymes that are involved in the degrada-
tion of proteins or phospholipids, such as cysteine proteinase and aspartic
proteinase. In addition to these genes, the expression of genes for glutathione
transferase in carnation (Meyer et al., 1991) and in-chain fatty acid hydroxy-
lase, fatty acid elongase, and allene oxide synthase in daylily (Panavas et al.,
1999) has been reported to increase during senescence.
In PCD of animal cells, caspase, a cysteine proteinase, plays a
very important role. Caspase has not been isolated in plants, but vacu-
olar processing enzyme (VPE) is a protease that exhibits caspase a­ ctivity
(­Hara-Nishimura et al., 2005). VPE is localized in the vacuoles and is
responsible for the maturation or activation of vacuolar proteins. Expression
of VPE homologs increases during petal senescence in Japanese morn-
ing glory (Yamada et al., 2009), carnation (Hoeberichts et al., 2007), and
­daffodil (Hunter et al., 2002).
The Bax (Bcl-2-associated X protein) inhibitor inhibits Bax activity
in the PCD of animal cells. Bax proteins have not been found in plants, but
genes for Bax inhibitor proteins have been isolated in several plant species.
In Japanese morning glory, the mRNA level of Bax inhibitor increases as


the petals senesce (Yamada et al., 2009). DAD1 (defender against apoptotic
cell death) is a protein that suppresses the progression of PCD in animal
cells. The mRNA level of a DAD1 homolog starts to decrease at early stages
of petal senescence in Alstroemeria (Breeze et al., 2004), while it decreases
at later stages in carnation (Hoeberichts et al., 2007), gladiolus (Yamada
et al., 2004), and Iris (Van Doorn et al., 2003). However, the role of DAD1 in
flower senescence is unknown.

4.4.3  Autophagy in Petal Senescence

Autophagy is often observed in plant cells at later stages of senescence.
Autophagy, or “self-eating,” is a conserved mechanism for the degradation
of cellular contents in eukaryotic species. This occurs by the uptake of cyto-
plasmic constituents into the vacuole, where they are degraded by vacuolar
hydrolases in plants (Bassham, 2007; Li and Vierstra, 2012). In senesc-
ing petals of morning glory (Ipomoea purpurea), numerous vesicles and
cytoplasmic components have been observed in the vacuoles by electron
microscopy, indicating autophagic processes (Matile and Winkenbach,
1971). Similar autophagic phenomena have been observed in senescing pet-
als of carnation (Smith et al., 1992).
Several genes involved in autophagic processes (autophagy-related
genes [ATGs]) have been identified in budding yeast (Saccharomyces
­cerevisiae) (Ohsumi, 2001), and ATG homologs have been isolated in
Arabidopsis thaliana (Bassham et al., 2006). mRNA levels for ATG4 and
ATG8 homologs in Japanese morning glory (Yamada et al., 2009; Shibuya
et al., 2009, 2011) and for ATG8 homologs in petunia increase during petal
senescence (Shibuya et al., 2013). The increase in mRNA levels of ATG
homologs may reflect the activation of autophagy; however, it is difficult to
accurately evaluate autophagic activity. Autophagy is proposed to play an
important role in nutrient recycling in leaf senescence (Bassham, 2007),
and this is likely the case for its role in petal senescence. Autophagy has
been thought to be closely associated with cell death in plants; however, it
is unclear whether autophagy is the direct cause of cell death during petal

4.5  Conclusions and Future Perspectives

Ethylene is clearly a key factor that regulates flower senescence in many
plant species. As ethylene biosynthetic and signaling pathways have been
well characterized, it is now possible to control the effects of ethylene by
chemical treatment or genetic modification. However, several issues in


flower senescence remain to be elucidated. In particular, the mechanisms

regulating ethylene-independent senescence are largely unknown.
It is unclear how differences in responses to ethylene among plant spe-
cies arise. In flowers that exhibit very low ethylene sensitivity, such as chrysan-
themum and gladiola, genes for ethylene receptors are expressed in the petals
(Arora et al., 2006; Narumi et al., 2005). Thus, ethylene should be perceived
and the signal transmitted in these flowers as in flowers with high ethylene
sensitivity. The ethylene signal somehow may not switch on downstream path-
ways that induce cell death in flowers that do not respond to ethylene.
Important roles of the gynoecia or stamen in the regulation of petal
senescence are highlighted in flowers with high ethylene sensitivity, such
as carnation and Portulaca (Ichimura and Suto, 1998; Shibuya et al., 2000).
However, the detailed mechanisms of interorgan regulation in natural
senescence of flowers remain to be elucidated.
In flowers that show ethylene-independent senescence, little is known
of the regulatory pathways of petal senescence. Since inhibitors of RNA and
protein synthesis delay senescence in these flowers, genes that actively reg-
ulate senescence should exist. Although many genes that show changes in
expression during senescence have been isolated, key regulators of flower
senescence have not been identified. A lack of good model plants that are
transformable makes it difficult to determine the function of isolated genes.
Recently, Japanese morning glory has been proposed as a model plant to
study ethylene-independent senescence; it is already used for functional
analysis (Shibuya, 2012).
Physiological and molecular biological studies on flower senescence
have made substantial contributions to the development of postharvest
technologies in cut flowers. However, the mechanisms behind flower senes-
cence are not fully understood. Further advances in the knowledge of
postharvest biology will help the development of innovative technology to
improve the postharvest quality of flowers.

Alonso, J.M., Hirayama, T., Roman, G., Nourizadeh, S., and Ecker, J.R. 1999.
EIN2, a bifunctional transducer of ethylene and stress responses in
Arabidopsis. Science 284, 2148–2152.
Arora, A., Watanabe, S., Ma, B., Takada, K., and Ezura, H. 2006. A novel eth-
ylene receptor homolog gene isolated from ethylene-insensitive flowers
of gladiolus (Gladiolus grandiflora Hort.). Biochemical and Biophysical
Research Communications 351, 739–744.
Bai, S.Y., Willard, B., Chapin, L.J., Kinter, M.T., Francis, D.M., Stead, A.D.,
and Jones, M.L. 2010. Proteomic analysis of pollination-induced corolla
senescence in petunia. Journal of Experimental Botany 61, 1089–1109.


Barden, L.E., and Hanan, J.J. 1972. Effect of ethylene on carnation keeping
life. Journal of the American Society for Horticultural Science 97, 785–788.
Bassham, D.C. 2007. Plant autophagy—more than a starvation response.
Current Opinion in Plant Biology 10, 587–593.
Bassham, D.C., Laporte, M., Marty, F., Moriyasu, Y., Ohsumi, Y., Olsen,
L.J., and Yoshimoto, K. 2006. Autophagy in development and stress
responses of plants. Autophagy 2, 2–11.
Borochov, A., Mayak, S., and Halevy, A.H. 1976a. Combined effects of abscisic
acid and sucrose on growth and senescence of rose flowers. Physiologia
Plantarum 36, 221–224.
Borochov, A., Tirosh, T., and Halevy, A.H. 1976b. Abscisic acid content of
senescing petals on cut rose flowers as affected by sucrose and water
stress. Plant Physiology 58, 175–178.
Breeze, E., Wagstaff, C., Harrison, E., Bramke, I., Rogers, H., Stead, A.,
Thomas, B., and Buchanan-Wollaston, V. 2004. Gene expression pat-
terns to define stages of postharvest senescence in Alstroemeria petals.
Plant Biotechnology Journal 2, 155–168.
Burg, S.P., and Dijkman, M.J. 1967. Ethylene and auxin participation in
pollen induded fading of Vanda orchid blossoms. Plant Physiology 42,
Celikel, F.G., and Reid, M.S. 2002. Postharvest handling of stock (Matthiola
incana). HortScience 37, 144–147.
Chang, C., Kwok, S.F., Bleecker, A.B., and Meyerowitz, E.M. 1993. Arabidopsis
ethylene-response gene ETR1: Similarity of product to two-component
regulators. Science 262, 539–544.
Chang, H., Jones, M.L., Banowetz, G.M., and Clark, D.G. 2003. Overproduction
of cytokinins in petunia flowers transformed with PSAG12-IPT delays
corolla senescence and decreases sensitivity to ethylene. Plant
Physiology 132, 2174–2183.
Chang, Y.S., and Chen, H.C. 2001. Variability between silver thiosulfate and
1-naphthaleneacetic acid applications in prolonging bract longevity of
potted bougainvillea. Scientia Horticulturae 87, 217–224.
Chao, Q., Rothenberg, M., Solano, R., Roman, G., Terzaghi, W., and Ecker, J.R.
1997. Activation of the ethylene gas response pathway in Arabidopsis by
the nuclear protein ETHYLENE-INSENSITIVE3 and related proteins.
Cell 89, 1133–1144.
Clark, D.G., Richards, C., Hilioti, Z., LindIversen, S., and Brown, K. 1997.
Effect of pollination on accumulation of ACC synthase and ACC oxidase
transcripts, ethylene production and flower petal abscission in gera-
nium (Pelargonium × hortorum L.H. Bailey). Plant Molecular Biology
34, 855–865.
Cook, D., Rasche, M., and Eisinger, W. 1985. Regulation of ethylene bio-
synthesis and action in cut carnation flower senescence by cytoki-
nins. Journal of the American Society for Horticultural Science 110,


Dervinis, C., Clark, D.G., Barrett, J.E., and Nell, T.A. 2000. Effect of polli-
nation and exogenous ethylene on accumulation of ETR1 homologue
transcripts during flower petal abscission in geranium (Pelargonium ×
hortorum L.H. Bailey). Plant Molecular Biology 42, 847–856.
Doi, M., Nakagawa, Y., Watabe, S., Aoe, K., Inamoto, K., and Imanishi,  H.
2003. Ethylene-induced leaf yellowing in cut chrysanthemums
(Dendranthema grandiflora Kitamura). Journal of the Japanese Society
for Horticultural Science 72, 533–535.
Eisinger, W. 1977. Role of cytokinins in carnation flower senescence. Plant
Physiology 59, 707–709.
Elgar, H.J., Woolf, A.B., and Bieleski, R.L. 1999. Ethylene production by three
lily species and their response to ethylene exposure. Postharvest Biology
and Technology 16, 257–267.
Eze, J.M.O., Mayak, S., Thompson, J.E., and Dumbroff, E.B. 1986. Senescence
in cut carnation flowers: temporal and physiological relationships
among water status, ethylene, abscisic acid and membrane permeability.
Physiologia Plantarum 68, 323–328.
Fromm, J., Hajirezaei, M., and Wilke, I. 1995. The biochemical response of
electrical signaling in the reproductive system of Hibiscus plants. Plant
Physiology 109, 375–384.
Gilissen, L.J.W. 1977. Style-controlled wilting of flower. Planta 133, 275–280.
Gilissen, L.J.W., and Hoekstra, F.A. 1984. Pollination-induced corolla wilting
in Petunia hybrida rapid transfer through the style of a wilting-inducing
substance. Plant Physiology 75, 496–498.
Goh, C.J., Halevy, A.H., Engel, R., and Kofranek, A.M. 1985. Ethylene evolution
and sensitivity in cut orchid flowers. Scientia Horticulturae 26, 57–67.
Goto, R., Aida, R., Shibata, M., and Ichimura, K. 1999. Role of ethylene
on flower senescence of Torenia. Journal of the Japanese Society for
Horticultural Science 68, 263–268.
Guo, H., and Ecker, J.R. 2003. Plant responses to ethylene gas are mediated
by SCF(EBF1/EBF2)-dependent proteolysis of EIN3 transcription
­factor. Cell 115, 667–677.
Halevy, A.H., Mayak, S., Tirosh, T., Spiegelstein, H., and Kofranek, A.M.
1974. Opposing effects of abscisic acid on senescence of rose flowers.
Plant and Cell Physiology 15, 813–821.
Hanley, K.M., and Bramlage, W.J. 1989. Endogenous levels of abscisic acid
in aging carnation flower parts. Journal of Plant Growth Regulation 8,
Hara-Nishimura, I., Hatsugai, N., Nakaune, S., Kuroyanagi, M., and
Nishimura, M. 2005. Vacuolar processing enzyme: An executor of plant
cell death. Current Opinion in Plant Biology 8, 404–408.
Henskens, J.A., Rouwendal, G.J., ten Have, A., and Woltering, E.J. 1994.
Molecular cloning of two different ACC synthase PCR fragments in
carnation flowers and organ-specific expression of the corresponding
genes. Plant Molecular Biology 26, 453–458.


Hiraya, T., Shimizu, H., and Ichimura, K. 2002. Effects of STS, 1-MCP
and sucrose on the vase life of cut Oxypetalum caeruleum flowers.
Horticulture Research (Japan), 1, 67–70 (in Japanese with English
Hoeberichts, F.A., Van Doorn, W.G., Vorst, O., Hall, R.D., and van Wordragen,
M.F. 2007. Sucrose prevents up-regulation of senescence-associated
genes in carnation petals. Journal of Experimental Botany 58, 2873–2885.
Hoekstra, F.A., and Weges, R. 1986. Lack of control by early pistillate ethylene
of the accelerated wilting of Petunia hybrida flowers. Plant Physiology 80,
Hopkins, M., Taylor, C., Liu, Z., Ma, F., McNamara, L., Wang, T.W., and
Thompson, J.E. 2007. Regulation and execution of molecular disassem-
bly and catabolism during senescence. New Phytology 175, 201–214.
Hua, J., and Meyerowitz, E.M. 1998. Ethylene responses are negatively regu-
lated by a receptor gene family in Arabidopsis thaliana. Cell 94, 261–271.
Hua, J., Chang, C., Sun, Q., and Meyerowitz, E.M. 1995. Ethylene insensitiv-
ity conferred by Arabidopsis ERS gene. Science 269, 1712–1714.
Huang, Y., Li, H., Hutchison, C.E., Laskey, J., and Kieber, J.J. 2003. Biochemical
and functional analysis of CTR1, a protein kinase that negatively regu-
lates ethylene signaling in Arabidopsis. Plant Journal 33, 221–233.
Hunter, D.A., Ferrante, A., Vernieri, P., and Reid, M.S. 2004a. Role of abscisic
acid in perianth senescence of daffodil (Narcissus pseudonarcissus
‘Dutch Master’). Physiologia Plantarum 121, 313–321.
Hunter, D.A., Steele, B.C., and Reid, M.S. 2002. Identification of genes associ-
ated with perianth senescence in daffodil (Narcissus pseudonarcissus L.
‘Dutch Master’). Plant Science 163, 13–21.
Hunter, D.A., Yi, M.F., Xu, X.J., and Reid, M.S. 2004b. Role of ethylene in
perianth senescence of daffodil (Narcissus pseudonarcissus L. ‘Dutch
Master’). Postharvest Biology and Technology 32, 269–280.
Ichimura, K. 2010. Postharvest physiology of cut flowers: progress and future
aspects. Bulletin of National Institute of Floriculture Sciences 10, 11–53
(in Japanese).
Ichimura, K., and Goto, R. 2000. Acceleration of senescence by pollination of
cut ‘Asuka-no-nami’ Eustoma flowers. Journal of the Japanese Society for
Horticultural Science 69, 166–170.
Ichimura, K., and Goto, R. 2002. Extension of vase life of cut Narcissus tazetta
var. chinensis flowers by combined treatment with STS and gibberellin A3.
Journal of the Japanese Society for Horticultural Science 71, 226–230.
Ichimura, K., and Suto, K. 1998. Role of ethylene in acceleration of flower
senescence by filament wounding in Portulaca hybrid. Physiologia
Plantarum 104, 603–607.
Ichimura, K., Kishimoto, M., Norikoshi, R., Kawabata, Y., and Yamada, K.
2005. Soluble carbohydrates and variation in vase-life of cut rose cultivars
‘Delilah’ and ‘Sonia’. Journal of Horticultural Science and Biotechnology
80, 280–286.


Ichimura, K., Niki, T., and Shimizu-Yumoto, H. 2007. Factors involved in the
vase life of cut snapdragon flowers. Horticulture Research (Japan) 6
(Suppl. 2), 341 (in Japanese).
Ichimura, K., Shimamura, M., and Hisamatsu, T. 1998. Role of ethylene in
senescence of cut Eustoma flowers. Postharvest Biology and Technology
14, 193–198.
Ichimura, K., Shimizu-Yumoto, H., and Goto, R. 2009a. Ethylene produc-
tion by gynoecium and receptacle is associated with sepal abscission
in cut Delphinium flowers. Postharvest Biology and Technology 52,
Ichimura, K., Yamada, T., Yoshida, S., Pun, U.K., Tanase, K., and Shimizu-
Yumoto, H. 2009b. Ethylene regulates programmed cell death
(PCD) associated with petal senescence in carnation flowers. Acta
Horticulturae 847, 185–190.
Iordachescu, M., and Verlinden, S. 2005. Transcriptional regulation of three
EIN3-like genes of carnation (Dianthus caryophyllus L. cv. Improved
White Sim) during flower development and upon wounding, polli-
nation,  and ethylene exposure. Journal of Experimental Botany 56,
Jones, M.L., and Woodson, W.R. 1997. Pollination-induced ethylene in carna-
tion: role of stylar ethylene in corolla senescence. Plant Physiology 115,
Jones, M.L., and Woodson, W.R. 1999a. Differential expression of three mem-
bers of the 1-aminocyclopropane-1-carboxylate synthase gene family in
carnation. Plant Physiology 119, 755–764.
Jones, M.L., and Woodson, W.R. 1999b. Inter-organ signaling following pol-
lination in carnations. Journal of the American Society for Horticultural
Science 124, 598–604.
Jones, R.B., Serek, M., Kuo, C.L., and Reid, M.S. 1994. The effect of protein
synthesis inhibition on petal senescence in cut bulb flowers. Journal of
the American Society for Horticultural Science 119, 1243–1247.
Kato, M., Shimizu, H., Onozaki, T., Tanikawa, N., Ikeda, H., Hisamatsu, T.,
and Ichimura, K. 2002. Role of ethylene in senescence of pollinated
and unpollinated Campanula medium flowers. Journal of the Japanese
Society for Horticultural Science 71, 385–387.
Kende, H. 1993. Ethylene biosynthesis. Annual Review of Plant Physiology
and Plant Molecular Biology 44, 283–307.
Kevany, B.M., Tieman, D.M., Taylor, M.G., Cin, V.D., and Klee, H.J. 2007.
Ethylene receptor degradation controls the timing of ripening in tomato
fruit. Plant Journal 51, 458–467.
Kieber, J.J., Rothenberg, M., Roman, G., Feldmann, K.A., and Ecker, J.R.
1993. CTR1, a negative regulator of the ethylene response pathway in
Arabidopsis, encodes a member of the raf family of protein kinases. Cell
72, 427–441.


Kosugi, Y., Shibuya, K., Tsuruno, N., Iwazaki, Y., Mochizuki, A., Yoshioka,
T., Hashiba, T., and Satoh, S. 2000. Expression of genes responsible for
ethylene production and wilting are differently regulated in carnation
(Dianthus caryophyllus L.) petals. Plant Science 158, 139–145.
Kuroda, S., Hakata, M., Hirose, Y., Shiraishi, M., and Abe, S. 2003. Ethylene
production and enhanced transcription of an ethylene receptor gene,
ERS1, in Delphinium during abscission of florets. Plant Physiology and
Biochemistry 41, 812–820.
Kuroda, S., Hirose, Y., Shiraishi, M., Davies, E., and Abe, S. 2004. Co-expression
of an ethylene receptor gene, ERS1, and ethylene signaling regulator
gene, CTR1, in Delphinium during abscission of florets. Plant Physiology
and Biochemistry 42, 745–751.
Langston, B.J., Bai, S., and Jones, M.L. 2005. Increases in DNA fragmenta-
tion and induction of a senescence-specific nuclease are delayed during
corolla senescence in ethylene-insensitive (etr1-1) transgenic petunias.
Journal of Experimental Botany 56, 15–23.
Larsen, P.B., Ashworth, E.N., Jones, M.L., and Woodson, W.R. 1995.
Pollination-induced ethylene in carnation: Role of pollen tube growth
and sexual compatibility. Plant Physiology 108, 1405–1412.
Li, F., and Vierstra, R.D. 2012. Autophagy: a multifaceted intracellular system
for bulk and selective recycling. Trends in Plant Science 17, 526–537.
Lin, Z., Zhong, S., and Grierson, D. 2009. Recent advances in ethylene
research. Journal of Experimental Botany 60, 3311–3336.
Lindstrom, J.T., Lei, C.H., Jones, M.L., and Woodson, W.R. 1999. Accumulation
of 1-aminocyclopropane-1-carboxylic acid (ACC) in petunia pollen is
associated with expression of a pollen-specific ACC synthase late in
development. Journal of the American Society for Horticultural Science
124, 145–151.
Lovell, P.J., Lovell, P.H., and Nichols, R. 1987. The control of flower senes-
cence in petunia (Petunia hybrida). Annals of Botany 60, 49–59.
Lurie, S. 1998. Postharvest heat treatments. Postharvest Biology and
Technology 14, 257–269.
Macnish, A.J., Jiang, C.Z., and Reid, M.S. 2010. Treatment with thidiazuron
improves opening and vase life of iris flowers. Postharvest Biology and
Technology 56, 77–84.
Matile, P., and Winkenbach, F. 1971. Function of lysosomes and lysosomal
enzymes in the senescing corolla of the morning glory (Ipomoea
­purpurea). Journal of Experimental Botany 22, 759–771.
Mayak, S., and Dilley, D.R. 1976a. Effect of sucrose on response of cut car-
nation to kinetin, ethylene, and abscisic acid. Journal of the American
Society for Horticultural Science 101, 583–585.
Mayak, S., and Dilley, D.R. 1976b. Regulation of senescence in carnation
(Dianthus caryophyllus): Effect of abscisic acid and carbon dioxide on
ethylene production. Plant Physiology 58, 663–665.
Mayak, S., and Halevy, A.H. 1970. Cytokinin activity in rose petals and its rela-
tion to senescence. Plant Physiology 46, 497–499.


Meyer Jr., R.C., Goldsbrough, P.B., and Woodson, W.R. 1991. An ethylene-
responsive flower senescence-related gene from carnation encodes
a protein homologous to glutathione s-transferases. Plant Molecular
Biology 17, 277–281.
Momonoi, K., Shoji, K., and Yoshida, K. 2007. Cloning and characterization of
ACC oxidase genes from tulip. Plant Biotechnology 24, 241–246.
Mor, Y., Halevy, A.H., Spiegelstein, H., and Mayak, S. 1985. The site of
1-­aminocyclopropane-1-carboxylic acid synthesis in senescing carna-
tion ­petals. Physiologia Plantarum 65, 196–202.
Mor, Y., Reid, M.S., and Kofranek, A.M. 1980. Role of the ovary in carnation
senescence. Scientia Horticulturae 13, 377–383.
Mor, Y., Reid, M.S., and Kofranek, A.M. 1984. Pulse treatments with silver
thiosulfate and sucrose improve the vase life of sweet peas. Journal of
the American Society for Horticultural Science 109, 866–868.
Mor, Y., Spiegelstein, H., and Halevy, A.H. 1983. Inhibition of ethylene
biosynthesis in carnation petals by cytokinin. Plant Physiology 71,
Müller, R., Lind-Iversen, S., Stummann, B.M., and Serek, M. 2000a. Expression
of genes for ethylene biosynthetic enzymes and an ethylene receptor
in senescing flowers of miniature potted roses. Journal of Horticultural
Science and Biotechnology 75, 12–18.
Müller, R., Owen, C.A., Xue, Z.T., Welander, M., and Stummann, B.M. 2002.
Characterization of two CTR-like protein kinases in Rosa hybrida and
their expression during flower senescence and in response to ethylene.
Journal of Experimental Botany 53, 1223–1225.
Müller, R., Owen, C.A., Xue, Z.T., Welander, M., and Stummann, B.M. 2003.
The transcription factor EIN3 is constitutively expressed in miniature
roses with differences in postharvest life. Journal of horticultural sci-
ence & biotechnology 78, 10–14.
Müller, R., Stummann, B.M., and Serek, M. 2000b. Characterization of an
ethylene receptor family with differential expression in rose (Rosa
­hybrida L.) flowers. Plant Cell Reports 19, 1232–1239.
Nadeau, J.A., Xian, S.Z., Nair, H., and O’neill, S.D. 1993. Temporal and spa-
tial regulation of 1-aminocyclopropane-1-carboxylate oxidase in the
p­ollination-induced senescence of orchid flowers. Plant Physiology 103,
Narumi, T., Kanno, Y., Suzuki, M., Kishimoto, S., Ohmiya, A., and Satoh, S.
2005. Cloning of a cDNA encoding an ethylene receptor (DG-ERS1)
from chrysanthemum and comparison of its mRNA level in ethylene-
sensitive and -insensitive cultivars. Postharvest Biology and Technology
36, 21–30.
Nichols, R. 1966. Ethylene production during senescence of flowers. Journal
of Horticultural Science and Biotechnology 41, 279–290.
Nichols, R. 1968. The response of carnations (Dianthus caryophyllus) to eth-
ylene. Journal of Horticultural Science 43, 335–349.


Norikoshi, R., Niki, T., and Ichimura, K. 2008. Changes in ACO activity and
two ACO gene expression in petals, styles and ovary during flower
senescence in cut carnations. Horticulture Research (Japan) 7 (Suppl. 2),
330 (in Japanese).
Ohsumi, Y. 2001. Molecular dissection of autophagy: two ubiquitin-like sys-
tems. National Review of Molecular Cell Biology 2, 211–216.
Okamoto, M., and Ichimura, K. 2010. Changes in ethylene production by pol-
lination in cut Delphinium grandiflorum. Horticulture Research (Japan)
9 (Suppl. 1), 218 (in Japanese).
O’Neill, S.D., Nadeau, J.A., Zhang, X.S., Bui, A.Q., and Halevy, A.H. 1993.
Interorgan regulation of ethylene biosynthetic genes by pollination.
Plant Cell 5, 419–432.
Onozaki, T., Ikeda, H., and Shibata, M. 2004. Video evaluation of ethylene
sensitivity after anthesis in carnation (Dianthus caryophyllus L.) flowers.
Scientia Horticulturae 99, 187–197.
Panavas, T., Pikula, A., Reid, P.D., Rubinstein, B., and Walker, E.L. 1999.
Identification of senescence-associated genes from daylily petals. Plant
Molecular Biology 40, 237–248.
Panavas, T., Walker, E.L., and Rubinstein, B. 1998. Possible involvement of
abscisic acid in senescence of daylily petals. Journal of Experimental
Botany 49, 1987–1997.
Park, K.Y., Drory, A., and Woodson, W.R. 1992. Molecular cloning of an
1-aminocyclopropane-1-carboxylate synthase from senescing carnation
flower petals. Plant Molecular Biology 18, 377–386.
Porat, R., and Halevy, A.H. 1993. Enhancement of petunia and dendrobium
flower senescence by jasmonic acid methyl ester is via the promotion of
ethylene production. Plant Growth Regulation 13, 297–301.
Porat, R., Borochov, A., and Halevy, A.H. 1994. Pollination-induced changes
in ethylene production and sensitivity to ethylene in cut dendrobium
orchid flowers. Scientia Horticulturae 58, 215–221.
Porat, R., Halevy, A.H., Serek, M., and Borochov, A. 1995a. An increase in
ethylene sensitivity following pollination is the initial event trigger-
ing an increase in ethylene production and enhanced senescence of
Phalaenopsis orchid flowers. Physiologia Plantarum 93, 778–784.
Porat, R., Reiss, N., Atzorn, R., Halevy, A.H., and Borochov, A. 1995b.
Examination of the possible involvement of lipoxygenase and jasmo-
nates in pollination-induced senescence of Phalaenopsis and Dendrobium
orchid flowers. Physiologia Plantarum 94, 205–210.
Qiao, H., Chang, K.N., Yazaki, J., and Ecker, J.R. 2009. Interplay between
­ethylene, ETP1/ETP2 F-box proteins, and degradation of EIN2 triggers
ethylene responses in Arabidopsis. Genes and Development 23, 512–521.
Qiao, H., Shen, Z.X., Huang, S.S.C., Schmitz, R.J., Urich, M.A., Briggs,
S.P., and Ecker, J.R. 2012. Processing and subcellular trafficking of
ER-tethered EIN2 control response to ethylene gas. Science 338,


Reid, M.S., Fujino, D.W., Hoffman, N.E., and Whitehead, C.S. 1984.
1-Aminocyclopropane-1-carboxylic acid (ACC): The transmitted
­stimulus  in pollinated flowers. Journal of Plant Growth Regulation 3,
Rogers, H.J. 2006. Programmed cell death in floral organs: How and why do
flowers die? Annals of Botany 97, 309–315.
Rogers, H.J. 2013. From models to ornamentals: How is flower senescence
regulated? Plant Molecular Biology 82, 563–574.
Sacalis, J.N., and Lee, J.S. 1987. Promotion of floral longevity by the ovary
in carnation flowers. Journal of the American Society for Horticultural
Science 112, 118–121.
Sacalis, J.N., and Nichols, R. 1980. Effects of 2,4-D uptake on petal senes-
cence in cut carnation flowers. HortScience 15, 499–500.
Sakai, H., Hua, J., Chen, Q.G., Chang, C., Medrano, L.J., Bleecker, A.B., and
Meyerowitz, E.M. 1998. ETR2 is an ETR1-like gene involved in ethylene
signaling in Arabidopsis. Proceedings of the National Academy of Sciences
USA 95, 5812–5817.
Saks, Y., Vanstaden, J., and Smith, M.T. 1992. Effect of gibberellic acid on car-
nation flower senescence: Evidence that the delay of carnation flower
senescence by gibberellic acid depends on the stage of flower develop-
ment. Plant Growth Regulation 11, 45–51.
Serek, M., Jones, R.B., and Reid, M.S. 1994. Role of ethylene in opening and
senescence of Gladiolus sp. flowers. Journal of the American Society for
Horticultural Science 119, 1014–1019.
Sexton, R., Laird, G., and Van Doorn, W.G. 2000. Lack of ethylene involve-
ment in tulip petal abscission. Physiologia Plantarum 108, 321–329.
Shibuya, K. 2012. Molecular mechanisms of petal senescence in ornamental
plants. Journal of the Japanese Society for Horticultural Science 81, 140–149.
Shibuya, K., and Clark, D.G. 2006. Ethylene: Current status and future direc-
tions of using transgenic techniques to improve flower longevity of
ornamental crops. Journal of Crop Improvement 18, 391–412.
Shibuya, K., and Ichimura, K. 2010. Depression of autocatalytic ethylene pro-
duction by high-temperature treatment in carnation flowers. Journal of
the Japanese Society for Horticultural Science 79, 97–102.
Shibuya, K., Barry, K.G., Ciardi, J.A., Loucas, H.M., Underwood, B.A.,
Nourizadeh, S., Ecker, J.R., Klee, H.J., and Clark, D.G. 2004. The central
role of PhEIN2 in ethylene responses throughout plant development in
petunia. Plant Physiology 136, 2900–2912.
Shibuya, K., Nagata, M., Tanikawa, N., Yoshioka, T., Hashiba, T., and
Satoh, S. 2002. Comparison of mRNA levels of three ethylene receptors
in senescing flowers of carnation (Dianthus caryophyllus L.). Journal of
Experimental Botany 53, 399–406.
Shibuya, K., Niki, T., and Ichimura, K. 2013. Pollination induces a­ utophagy
in petunia petals via ethylene. Journal of Experimental Botany 64,


Shibuya, K., Shimizu, K., Yamada, T., and Ichimura, K. 2011. Expression of
autophagy-associated ATG8 genes during petal senescence in Japanese
morning glory. Journal of the Japanese Society for Horticultural Science
80, 89–95.
Shibuya, K., Yamada, T., Suzuki, T., Shimizu, K., and Ichimura, K. 2009.
InPSR26, a putative membrane protein, regulates programmed cell
death during petal senescence in Japanese morning glory. Plant
Physiology 149, 816–824.
Shibuya, K., Yoshioka, T., Hashiba, T., and Satoh, S. 2000. Role of the gynoe-
cium in natural senescence of carnation (Dianthus caryophyllus L.)
­flowers. Journal of Experimental Botany 51, 2067–2073.
Shimizu, H., and Ichimura, K. 2001. Involvement of ethylene in accelera-
tion of flower senescence by filament touching in Portulaca hybrid.
Horticulture Research (Japan) 70 (Suppl. 2), 467 (in Japanese).
Shimizu-Yumoto, H., and Ichimura, K. 2006a. Involvement of high sensitivity of
flag petal to ethylene on its closing in cut sweet pea flowers. Journal of the
Japanese Society for Horticultural Science 75 (Suppl. 1), 246 (in Japanese).
Shimizu-Yumoto, H., and Ichimura, K. 2006b. Senescence of Eustoma flowers
as affected by pollinated area of the stigmatic surface. Journal of the
Japanese Society for Horticultural Science 75, 66–71.
Shimizu-Yumoto, H., and Ichimura, K. 2009a. Abscisic acid, in combination
with sucrose, is effective as a pulse treatment to suppress leaf dam-
age and extend foliage vase-life in cut Eustoma flowers. Journal of
Horticultural Science and Biotechnology 84, 107–111.
Shimizu-Yumoto, H., and Ichimura, K. 2009b. Cultivar variation in the vase life
of unpollinated Eustoma flowers associated with ethylene. Horticulture
Research (Japan) 8, 359–364 (in Japanese with English abstract).
Singh, A., Evensen, K.B., and Kao, T.H. 1992. Ethylene synthesis and flo-
ral senescence following compatible and incompatible pollinations in
Petunia inflata. Plant Physiology 99, 38–45.
Smith, M.T., Saks, Y., and van Staden, J. 1992. Ultrastructural changes in the
petals of senescing flowers of Dianthus caryophyllus L. Annals of Botany
69, 277–285.
Solano, R., Stepanova, A., Chao, Q., and Ecker, J.R. 1998. Nuclear events in
ethylene signaling: a transcriptional cascade mediated by ETHYLENE-
Development 12, 3703–3714.
Song, L.L., and Peng, Y.H. 2004. Effect of cold storage on sensitivity of cut
lily to ethylene. Journal of Horticultural Science and Biotechnology 79,
Spanjers, A.W. 1981. Bioelectric potential changes in the style of Lilium lon-
giflorum Thunb. after self- and cross-pollination of the stigma. Planta
153, 1–5.
Stead, A.D. 1992. Pollination-induced flower senescence—A review. Plant
Growth Regulation 11, 13–20.


Stead, A.D., and Moore, K.G. 1979. Studies on flower longevity in Digitalis:
Pollination induced corolla abscission in Digitalis flowers. Planta 146,
Tanase, K., and Ichimura, K. 2006. Expression of ethylene receptors
Dl-ERS1-3 and Dl-ERS2, and ethylene response during flower senes-
cence in Delphinium. Journal of Plant Physiology 163, 1159–1166.
Tang, X., Gomes, A., Bhatia, A., and Woodson, W.R. 1994. Pistil-specific and
ethylene-regulated expression of 1-aminocyclopropane-1-carboxylate
oxidase genes in petunia flowers. Plant Cell 6, 1227–1239.
Tang, X.Y., Wang, H., Brandt, A.S., and Woodson, W.R. 1993. Organization
and structure of the 1-aminocyclopropane-1-carboxylate oxidase gene
family from Petunia hybrida. Plant Molecular Biology 23, 1151–1164.
ten Have, A., and Woltering, E.J. 1997. Ethylene biosynthetic genes are
­differentially expressed during carnation (Dianthus caryophyllus L.)
flower senescence. Plant Molecular Biology 34, 89–97.
Tripathi, S.K., and Tuteja, N. 2007. Integrated signaling in flower senescence:
an overview. Plant Signaling and Behaviour 2, 437–445.
Van Doorn, W.G., and Woltering, E.J. 2008. Physiology and molecular biology
of petal senescence. Journal of Experimental Botany 59, 453–480.
Van Doorn, W.G., Balk, P.A., van Houwelingen, A.M., Hoeberichts, F.A., Hall,
R.D., Vorst, O., van der Schoot, C., and van Wordragen, M.F. 2003.
Gene expression during anthesis and senescence in Iris flowers. Plant
Molecular Biology 53, 845–863.
van Staden, J., Featonbysmith, B.C., Mayak, S., Spiegelstein, H., and Halevy,
A.H. 1987. Cytokinins in cut carnation flowers. II. Relationship between
endogenous ethylene and cytokinin levels in the petals. Plant Growth
Regulation 5, 75–86.
Wagstaff, C., Chanasut, U., Harren, F.J.M., Laarhoven, L.J., Thomas, B.,
Rogers, H.J., and Stead, A.D. 2005. Ethylene and flower longevity
in Alstroemeria: relationship between petal senescence, abscis-
sion and ethylene biosynthesis. Journal of Experimental Botany 56,
Waki, K., Shibuya, K., Yoshioka, T., Hashiba, T., and Satoh, S. 2001. Cloning
of a cDNA encoding EIN3-like protein (DC-EIL1) and decrease in its
mRNA level during senescence in carnation flower tissues. Journal of
Experimental Botany 52, 377–379.
Wang, D., Fan, J., and Ranu, R.S. 2004. Cloning and expression of
aminocyclopropane-1-carboxylate synthase cDNA from rosa
(Rosa × ­hybrida). Plant Cell Reports 22, 422–429.
Wang, H., and Woodson, W.R. 1991. A flower senescence-related mRNA from
carnation shares sequence similarity with fruit ripening-related mRNAs
involved in ethylene biosynthesis. Plant Physiology 96, 1000–1001.
Wang, T.W., and Arteca, R.N. 1995. Identification and characterization of
cDNAs encoding ethylene biosynthetic enzymes from Pelargonium ×
hortorum cv Snow Mass leaves. Plant Physiology 109, 627–636.


Wang, Y., and Kumar, P.P. 2007. Characterization of two ethylene receptors
PhERS1 and PhETR2 from petunia: PhETR2 regulates timing of anther
dehiscence. Journal of Experimental Botany 58, 533–544.
Whitehead, C.S., Halevy, A.H., and Reid, M.S. 1984. Roles of ethylene and
1-aminocyclopropane-1-carboxylic acid in pollination and woundin-
duced senescence of Petunia hybrida flowers. Physiologia Plantarum 61,
Woltering, E.J., and Van Doorn, W.G. 1988. Role of ethylene in senescence
of petals—Morphological and taxonomical relationships. Journal of
Experimental Botany 39, 1605–1616.
Woltering, E.J., Balk, P.A., Nijenhuis-de Vries, M.A., Faivre, M., Ruys, G.,
Somhorst, D., Philosoph-Hadas, S., and Friedman, H. 2005. An auxin-
responsive 1-aminocyclopropane-1-carboxylate synthase is responsible
for differential ethylene production in gravistimulated Antirrhinum
majus L. flower stems. Planta 220, 403–413.
Woltering, E.J., deVrije, T., Harren, F., and Hoekstra, F.A. 1997. Pollination
and stigma wounding: Same response, different signal? Journal of
Experimental Botany 48, 1027–1033.
Woltering, E.J., Somhorst, D., and Vanderveer, P. 1995. The role of ethylene
in interorgan signaling during flower senescence. Plant Physiology 109,
Woodson, W.R., and Brandt, A.S. 1991. Role of the gynoecium in cytokinin-
induced carnation petal senescence. Journal of the American Society for
Horticultural Science 116, 676–679.
Woodson, W.R., Park, K.Y., Drory, A., Larsen, P.B., and Wang, H. 1992.
Expression of ethylene biosynthetic pathway transcripts in senescing
carnation flowers. Plant Physiology 99, 526–532.
Wu, M.J., Zacarias, L., and Reid, M.S. 1991. Variation in the senescence of car-
nation (Dianthus caryophyllus L.) cultivars. II. Comparison of sensitivity
to exogenous ethylene and of ethylene binding. Scientia Horticulturae
48, 109–116.
Wulster, G., Sacalis, J., and Janes, H.W. 1982. Senescence in isolated carnation
petals. Effects of indoleacetic acid and inhibitors of protein ­synthesis.
Plant Physiology 70, 1039–1043.
Xu, X.J., Gookin, T., Jiang, C.Z., and Reid, M. 2007. Genes associated with open-
ing and senescence of Mirabilis jalapa flowers. Journal of Experimental
Botany 58, 2193–2201.
Xu, Y., and Hanson, M.R. 2000. Programmed cell death during pollination-
induced petal senescence in petunia. Plant Physiology 122, 1323–1333.
Xue, J.Q., Li, Y.H., Tan, H., Yang, F., Ma, N., and Gao, J.P. 2008. Expression
of ethylene biosynthetic and receptor genes in rose floral tissues during
ethylene-enhanced flower opening. Journal of Experimental Botany 59,
Yamada, T., Ichimura, K., Kanekatsu, M., and Van Doorn, W.G. 2007a. Gene
expression in opening and senescing petals of morning glory (Ipomoea
nil) flowers. Plant Cell Reports 26, 823–835.


Yamada, T., Ichimura, K., and Van Doorn, W.G. 2007b. Relationship between
petal abscission and programmed cell death in Prunus yedoensis and
Delphinium belladonna. Planta 226, 1195–1205.
Yamada, T., Ichimura, K., Kanekatsu, M., and Van Doorn, W.G. 2009.
Homologues of genes associated with programmed cell death in a­ nimal
cells are differentially expressed during senescence of Ipomoea nil
­petals. Plant and Cell Physiology 50, 610–625.
Yamada, T., Takatsu, Y., Kasumi, M., Ichimura, K., and Van Doorn, W.G. 2006.
Nuclear fragmentation and DNA degradation during programmed cell
death in petals of morning glory (Ipomoea nil). Planta 224, 1279–1290.
Yamada, T., Takatsu, Y., Kasumi, M., Marubashi, W., and Ichimura, K. 2004.
A homolog of the defender against apoptotic death gene (DAD1) in
senescing gladiolus petals is down-regulated prior to the onset of pro-
grammed cell death. Journal of Plant Physiology 161, 1281–1283.
Yamane, K., and Ogata, R. 1995. Effects of cycloheximide on physiological
parameters of gladiolus florets during growth and senescence. Journal
of the Japanese Society for Horticultural Science 64, 411–416.
Yanagisawa, S., Yoo, S.D., and Sheen, J. 2003. Differential regulation of
EIN3 stability by glucose and ethylene signalling in plants. Nature 425,
Yang, S.F., and Hoffman, N.E. 1984. Ethylene biosynthesis and its regulation
in higher plants. Annual Review of Plant Physiology and Plant Molecular
Biology 35, 155–189.
Yangkhamman, P., Fukai, S., and Ichimura, K. 2005. Ethylene production and
vase life of cut carnation flowers under high temperature conditions.
Journal of the Japanese Society for Horticultural Science 74, 337–341.
Yangkhamman, P., Tanase, K., Ichimura, K., and Fukai, S. 2007. Depression
of enzyme activities and gene expression of ACC synthase and ACC oxi-
dase in cut carnation flowers under high-temperature conditions. Plant
Growth Regulation 53, 155–162.

Chapter 5

Mikal E. Saltveit
University of California, Davis, California

Abstract 140
5.1 Introduction 140
5.2 Why Measure Respiration? 141
5.3 Major Components of Respiration 142
5.3.1 Glycolysis 142
5.3.2 Pentose-Phosphate Shunt 142
5.3.3 Anaerobic Diversion 143
5.3.4 Tricarboxylic Acid Cycle 143
5.3.5 Electron Transport (Chemiosmotic
Phosphorylation) 145
5.4 Measurement of Respiration 146
5.4.1 Loss of Substrate, Heat Production,
and Water 146
5.4.2 Consumption of Oxygen and Production
of Carbon Dioxide 147 Static System 147 Flow-Through or Dynamic System 149
5.5 Sampling and Analyzing 151
5.6 Instruments and Techniques 152


5.7 Pre- and Postharvest Factors Affecting Respiration 152

5.7.1 Temperature Effects 152
5.7.2 Respiratory Quotient 154
5.7.3 Physical Stress 154
5.7.4 Internal Factors 155 Genotype 155 Type of Plant Part 155 Respiratory Climacteric 155
5.8 Conclusions and Future Perspectives 156
References 156

The three groups of catabolic reactions comprising respiration provide the
energy, the reducing power and carbon fragments used in subsequent ana-
bolic metabolic reactions. Horticultural crops are harvested either at the
optimal stage of maturity and quality (e.g., cucumber, lettuce, zucchini) or
at a stage of maturity that requires subsequent changes to reach maximal
quality (e.g., avocadoes, bananas, tomatoes). In the first group, respiratory
metabolism is suppressed in the postharvest environment to levels only
needed to maintain product quality. In the latter group, the often profound
changes that accompany ripening (e.g., pigment destruction and synthesis,
synthesis of characteristic aroma and flavors, tissue softening, and textural
changes) require significant inputs of energy and substrates derived from
respiration. Measurements of respiration (e.g., carbon dioxide production,
oxygen consumption) give a good indication of the metabolic activity of
the tissue, and thereby the rate of changes in product quality. The rate of
respiration and metabolic activity is tightly coupled to tissue temperature.
Both are reduced by lower product temperatures and increased by elevated
product temperatures. Some commodities (especially those of tropical and
subtropical origin, e.g., avocado, banana, bean, and squash) suffer physi-
ological damage (i.e., chilling injury) if exposed to temperatures below
10°C (50°F) for commodity-specific inductive periods. Respiratory and
metabolic activity can also be reduced by limiting the availability of oxygen
to the tissue. However, too little oxygen induces anaerobic respiration with
production of uncharacteristic compounds that can reduce product quality.

5.1 Introduction
The function of respiratory metabolism is threefold: to extract energy
stored in the chemical bonds of substrate molecules (e.g., glucose, fatty
acids, proteins) and convert it into high-energy phosphate bonds (i.e., ATP),

Respir ato ry Me ta bo lism

to capture reducing power (e.g., in NADH), and to produce small carbon

fragments from which more complex molecules can be synthesized. In
the presence of adequate levels of oxygen, the three sequential respiratory
reactions are glycolysis (i.e., the lysing of glucose to produce pyruvate), the
tricarboxylic acid cycle (i.e., the decarboxylation of pyruvate to CO2 and
production of high-energy compounds, e.g., NADH), and electron trans-
port (i.e., generation of ATP from high-energy compounds by chemiosmotic
phosphorylation). An offshoot of glycolysis, the pentose-phosphate shunt
diverts and reconfigures intermediates from glycolysis into substrates for
subsequent synthetic reactions (e.g., produces the 5-carbon ribose sugars
used in RNA and DNA synthesis). In the absence of oxygen, NAD+ regener-
ated during anaerobic respiration allows glycolysis to continue to extract a
portion of the energy contained in the glucose molecule.
Biological energy is stored and released through redox reactions.
Respiratory metabolism involves the transfer of energy in a substrate mol-
ecule (e.g., glucose) through the extraction and movement of electrons.
Redox (shorthand for reduction–oxidation reactions) involves a coupled
reaction where electrons are gained by a molecule (i.e., it is reduced) while
another molecule is oxidized (i.e., it loses electrons). In aerobic respira-
tion (i.e., in the presence of adequate oxygen), glucose is oxidized (i.e., it
furnishes high-energy electrons) and some of the energy in the electrons
is extracted in the production of ATP as they pass through a number of
enzymatically controlled steps, until they finally reduce oxygen to water
(i.e., transfer the electrons, with the accompanying protons, to oxygen). In
contrast to respiration, photosynthesis reduces carbon dioxide (i.e., adds
electrons to CO2) and oxidizes water (i.e., removes electrons from H 2O) to
produce glucose and oxygen (C6H12O6 and 6O2).
Carbohydrates (i.e., simple sugars like glucose derived from stored
starch) are the main respiratory substrate for most postharvest commodi-
ties. Although glucose contains 686 kcal/mol in chemical bonds, only
673 kcal/mol is available to do “work” because 13 kcal/mol is lost due to the
increase in disorder of the products (i.e., an increase in entropy). However,
only 281 kcal/mol of the 673 kcal/mol (42%) is captured in the 38 ATP
(7.4 kcal/mol) formed because of the inefficiencies of phosphorylation.

5.2  Why Measure Respiration?

Because respiration is tightly controlled so that no more ATP is produced
than is needed for metabolism, and because ATP production is tightly cou-
pled to respiration, the rate of respiration is normally a very accurate indica-
tor of the metabolic activity of the tissue. We do not measure respiration to
find only the rate of CO2 production or O2 consumption, but also the under-
lying rate of metabolism in the tissue. The rate of metabolism is usually


inversely related to the shelf life of the commodity. The higher the rate of
respiration, the more quickly there will be a decrease in the quality, nutri-
tion, or taste of the commodity. Since we cannot easily measure changes in
these parameters of shelf life, we use measurements of the rate of respira-
tion as their surrogate.

5.3  Major Components of Respiration

5.3.1 Glycolysis
As its name implies, the 10 enzymatic reactions in glycolysis break (or lyse)
the 6-carbon glucose molecule into two 3-carbon pyruvate molecules. The
reactions of glycolysis take place in the cytoplasm. Glucose is first phosphory-
lated in a series of reactions by two ATPs to produce fructose-1,6-diphosphate.
Living cells are homeostatic over short periods of time and tend to maintain
the status quo by carefully regulating the concentration of ATP. A key enzyme
in glycolysis is phosphofructokinase (PFK), a multicomponent enzyme that
converts fructose-6-phosphate into fructose-1,6-diphosphate, thereby pre-
paring the hexose for cleavage into two dissimilar phosphate-containing
3-carbon molecules (i.e., dihydroxyacetone phosphate and glyceraldehyde-
3-phosphate [G3P]), which are interconvertible. The rate of glycolysis is reg-
ulated primarily through controlling the activity of PFK, an enzyme that is
inhibited by ATP, one product of respiration. However, almost every enzyme
in the respiratory pathway is under some sort of metabolic control, indicating
the importance of careful regulation of respiration and metabolism.
Each molecule of G3P undergoes a series of reactions during which
one molecule of NADH and two molecules of ATP are produced. The result-
ing two molecules of the 3-carbon pyruvate contain all six carbons that were
in the original molecule of glucose. Two ATPs were needed to initiate the
reaction, but four ATPs and two NADHs were produced, for a net gain of
two ATPs and two NADHs. So while there are the same numbers of carbon
atoms in the two pyruvate molecules as there were in the original glucose
molecule, the pyruvate molecules have lost about 20% of the extractable
energy originally present in the glucose molecule. The remaining 80% is
extracted by the two subsequent reactions of respiration. Notice that no oxy-
gen was consumed, and no carbon dioxide was produced during glycolysis.

5.3.2  Pentose-Phosphate Shunt

Glucose-6-phosphate can be diverted from glycolysis to another oxidative
pathway that generates NADPH and five carbon substrates for further bio-
synthesis (e.g., ribose-5-phosphate for RNA synthesis).

Respir ato ry Me ta bo lism

There are two similar forms of nicotinamide adenine dinucleotide:

one with a phosphate group (NADPH) and one without a phosphate group
(NADH). In general, NADH participates in catabolic reactions, that is, reac-
tions of cellular respiration that break down molecules to release energy,
while NADPH participates in anabolic reactions, such as carbon fixation
during photosynthesis, that is, reactions that consume energy in order to
build up or synthesize larger molecules.

5.3.3  Anaerobic Diversion

NAD+ is needed for glycolysis to function. In the absence of oxygen (i.e.,
anaerobic), NAD+ accumulates in its reduced state (NADH) and all energy
production ceases with the subsequent death of the cell unless the oxidized
form (NAD+) can be regenerated. Anaerobic respiration starts with the
enzyme pyruvate decarboxylase removing a CO2 from pyruvate, forming
the 2-carbon acetaldehyde. However, acetaldehyde is a very reactive mol-
ecule and is quickly converted to ethanol by the enzyme alcohol dehydroge-
nase with the concomitant regeneration of NAD+. In some cells, especially
those that are less acidic, the enzyme lactate dehydrogenase regenerates
NAD+ during the production of lactic acid. However, the accumulation of
lactic acid acidifies the cell and stimulates the activity of pyruvate decar-
boxylase, which thereby shifts carbon flow from the production of lactate
to the production of ethanol. Differences in these two pathways are illus-
trated by the production of yogurt and beer. Lactic acid production during
fermentation to produce yogurt occurs without the release of CO2, while
production of ethanol during beer production is accompanied by the release
of CO2, which generates the effervescent beverage.

5.3.4  Tricarboxylic Acid Cycle

Pyruvate produced by glycolysis can enter a cyclic series of eight reactions
called the tricarboxylic acid cycle (TCA) (also called the citric acid cycle or
Krebs cycle) (Figure 5.1). Hans A. Krebs was the scientist who first eluci-
dated the cycle, and the first molecule in the cycle is citric acid, a tricarbox-
ylic acid. These reactions take place in the liquid portion (i.e., the matrix)
of the mitochondria, a semiautonomous organelle within eukaryotic cells.
Pyruvate is decarboxylated, yielding a molecule of CO2 and a 2-­carbon
acetate residue that is attached to a co-enzyme forming acetyl-CoA. The
2-carbon acetate group is then transferred to the 4-carbon oxaloacetate
molecule, forming the 6-carbon citric acid. As the cycle progresses, an
additional two molecules of CO2 are released, and the resulting 4-carbon
molecule is rearranged into the starting product of oxaloacetate, thereby


Proteins Fats
Simple sugar
(e.g., glucose)
Amino acids Fatty acids



Acetyl CoA

acid CO2

Reducing Power
3 O2 (NADH, FADH2)

Electron transport

3 H2O Mitochondria Cytoplasm

Figure 5.1  Overview of important features of cellular respiration.

completing the cycle. During the cycle, three NADHs, one FADH2, and one
ATP are produced from each pyruvate molecule (Figure 5.2).
High-energy molecules are not the only product of the TCA cycle. The
intermediates within the cycle can be substrates for subsequent synthetic
reactions. For example, pyruvate is used in the synthesis of acetaldehyde,
ethanol, lactic acid, and alanine; acetyl CoA is used in the synthesis of fatty
acids, cuticular compounds, isoprenoids, carotenoids, sterols, terpenes,
and aromatic amino acids; α-ketoglutarate is used in the synthesis of glu-
tamic acid, other amino acids, chlorophyll, cytochromes, and phytochrome;

Respir ato ry Me ta bo lism

Pyruvic acid
COOH Pyruvic
C=O decarboxylase CO2
Lactate Acetaldehyde
dehydrogenase Pyruvic H-C=O
2H+ oxidase CH3 NADH
CO2 dehydrogenase
Acetyl-CoA CH3
Lactic acid

To TCA cycle

Figure 5.2  The fate of pyruvic acid in respiratory metabolism.

and oxaloacetic acid is used in the synthesis of aspartic acid, alkaloids, and
nucleic acids.
Notice that in the TCA cycle, all the carbon from the glucose molecule
was released as CO2, but no oxygen has yet been consumed.

5.3.5  Electron Transport (Chemiosmotic Phosphorylation)

The preceding respiratory reactions have converted all the carbon in the
original glucose molecule into CO2, but in none of the previous reactions has
oxygen been consumed. The final reaction of respiratory metabolism regen-
erates NAD+ and converts the energy contained in NADH into ATP. During
this process, the electrons contained in NADH and FADH2 are transferred
to oxygen, the terminal electron acceptor, thereby producing water.
It was previously thought that the series of reactions generating ATP
from NADH (and FADH2) in the mitochondria were a series of coupled enzy-
matic reactions, with the product of one reaction furnishing the substrate
for the next reaction. It is now known that the production of ATP involves
using the high-energy electrons from NADH to pump protons across the
inner mitochondrial membrane, thereby establishing a proton gradient
between the intermembrane space and the matrix. The protons reenter the
matrix by passing through an H+ -ATPase complex embedded in the inner


membrane. This ATPase uses the energy in the returning protons to pro-
duce ATP from ADP and Pi (inorganic phosphate), a process called chemi-
osmotic phosphorylation.

5.4  Measurement of Respiration

Since the rate of respiration is so tightly coupled to the rate of cell metabo-
lism, measurements of respiration can afford an easy, nondestructive means
of monitoring the metabolic and physiological state of the tissues. Nowhere
in plant science is this truer than in the area of postharvest physiology,
where the events of senescence and ripening are often signaled by abrupt
changes in respiratory behavior. For this reason, postharvest physiologists
have spent considerable time devising means to measure respiration.
The overall reactions of respiratory metabolism consume substrate
molecules (i.e., carbohydrates, fatty acids, proteins, and oxygen) and pro-
duce high-energy compounds, CO2, and water. The rate of respiration could
be measured by the rate at which the substrates disappear or the products
appear. Apart from the water produced by respiration, which is relatively
trivial compared to the very high water content of plant (and particularly
fruit) tissues, all the substrates and products of respiration have been used
in determining respiration rate. They are: loss of substrate, loss of oxygen,
increase in carbon dioxide, and production of heat.
C6H12O6 + 6O2 → 6CO2 + 6H 2O + 38 ATP + 392 kcal (heat)

5.4.1  Loss of Substrate, Heat Production, and Water

It is theoretically possible to monitor the respiration of stored commodities
by measuring the loss of dry matter. During extended storage periods at
warm temperatures, the loss of dry matter of a respiring commodity can
be substantial. When glucose (a hexose sugar) is the substrate, 180 g of
glucose is lost for each 264 g of CO2 produced by the commodity. The rate
of dry weight loss (i.e., loss of glucose) can be estimated by multiplying
the weight of CO2 produced by 0.68: the ratio of the molecular weight of
glucose (180) divided by the molecular weight of the six CO2 produced
(264 = 6 × 44).
In addition to actual dry weight loss (which could be an important
consideration, for example, in onions being stored prior to dehydration), the
oxidation of substrates can be an important postharvest loss because of the
reduction in food value of the commodity to the consumer, reduced taste
quality (sweetness), and potential for acceleration of senescence in tissues
whose storage reserves have been depleted.

Respir ato ry Me ta bo lism

The rate of respiration has occasionally been measured by determin-

ing the rate at which heat is produced by the commodity. All of the heat
produced during respiration is lost from the tissue either by conduction to
the cold storage room atmosphere or by vaporization of water in the tissue.
If all the energy not captured in ATP (673 − 281 = 392 kcal) was lost from
the tissue by vaporizing water, 676 g of H 2O would be lost for each 180 g of
glucose oxidized (392 kcal/580 cal/g H 2O). Since 108 g of H 2O is produced
during respiration, there would be a net loss of 568 g of H 2O. This loss will
occur even in a water-saturated atmosphere. Apples at 0°C have a respira-
tion rate of about 4 mg CO2/kg-h, so 10 kg of apples would respire 1.2 g of
CO2, or about 0.82 g of glucose would be lost in a month (30 days). During
this time, 1.78 kcal of heat would have been produced [(1.2/264) × 392] and
could have led to the loss of 3.1 g of water (1.78 kcal/580 kcal/g H 2O).
Measurement of heat production is also important when engineer-
ing refrigeration systems used during postharvest handling and storage.
Calculation of heat production from the respiration equation shows that pro-
duction of 1 mg of CO2 yields 2.55 cal. In the language of the refrigeration
­engineer, a respiration rate of 1 mg CO2/kg-h indicates a heat production of
61.2 kcal/metric ton/day (220 BTU/tonne of produce/day). The British ther-
mal unit (BTU) is defined as the heat required to raise 1 lb of water by 1°F.

5.4.2  Consumption of Oxygen and

Production of Carbon Dioxide
Almost all practical methods for measuring the respiration of plant tis-
sues involve the estimation of uptake of O2 or release of CO2 by the tissue.
These methods are generally simple and have the great advantage of being
nondestructive. Indeed, they have often been used on commodities still
attached to the plant. Two general systems have been used for measuring
gas exchange: the static and the flow-through system.
In either system, respiration rates are usually expressed as weight
or volume of gas produced or consumed per kilogram of fresh weight of
product per hour, for example, 125 mg CO2/kg fresh wt/h or 25 ml O2/kg
fresh weight/h.  Static System

In the static system, tissue is placed in a sealed container and the accumu-
lation of CO2 or the depletion of O2 in the atmosphere is periodically mea-
sured (Figure 5.3). This method can be employed over brief periods of time,
especially with even very small portions of tissue (e.g., meristems, florets,
buds), but it has the disadvantage of being a nonequilibrium system. The
depletion of O2 and accumulation of CO2 or other gases (particularly C2H4)



O2 CO2

Carbon dioxide %



0 1 2 3 4 5
(a) (b)

Figure 5.3  Representation of the setup for measuring O2 consumption or

CO2 production in a static or closed system. (a) Depiction of a commodity in
a closed container. (b) Graph showing the increase in CO2 that would occur
as the commodity respired in a closed container. Note the derivation from
linearity as the CO2 concentration exceeds around 0.2%.

may strongly affect the tissue and its respiration rate. As can be seen in the
graph, the accumulation of CO2 becomes nonlinear at around 0.2%.
Problems with the accumulation of physiologically active concen-
trations of CO2 and C 2H4 can be overcome by including KOH to absorb
CO2 and KMnO 4 to oxidize C 2H4 . Indeed, some respirometers ( jargon for
a machine that measures respiration) measure the pressure (or volume)
change resulting from absorption of respired CO2 . The most notable of
these, the Warburg respirometer, was instrumental in determining the
pathway of glycolysis and has driven many postharvest physiologists to
strong drink because of the technical difficulties often encountered in
its use.
To calculate the rate of respiration in a static system, you need to know
the following: volume of container, weight of tissue, initial carbon dioxide
concentration, length of time, and final carbon dioxide concentration.
For example, assume that a 280 g apple is enclosed in a 1560 ml
container for 15 min. During that time, the CO2 concentration in the jar
increased from 0.03% to 0.23%, or the CO2 concentration increased 0.2%.
Converting the weight and time units to kilograms and hours gives a
weight of 0.28 kg and a sampling period of 15/60 = 0.25 h. The simplest
way to calculate the amount of CO2 produced during the sampling period
would be to multiply the difference in CO2 concentrations between the
beginning and end of the sampling period (i.e., 0.2%) by the total volume of
the container (0.2% × 1560 ml = 3.12 ml). Dividing that volume of CO2 by

Respir ato ry Me ta bo lism

the weight (in kg) and the sampling period (in hours) would give the rate
of CO2 production (i.e., 3.12 ml/(0.28 kg × 0.25 h) = 44.6 ml CO2/kg-h).
However, this calculation could be criticized because many assump-
tions were made that can significantly alter the answer. For example, it did
not take into account the fact that the volume occupied by the apple reduced
the total gas volume in the jar; i.e., the density of the tissue could deviate
from 1 and affect its volume, and the solubility of the gas being measured
in the cellular solution. The density of an apple is around 0.8 g/ml, so the
volume occupied by the apple would be 280/0.8 = 350 ml. The liquid in the
apple fruit will absorb some of the CO2 produced. The solubility of 100% CO2
in water at 20°C is 0.878 ml CO2/ml water (and increases with decreasing
temperatures). Taking all these caveats into consideration, we can conclude
that under normal physiological conditions, the rate of CO2 production can
be calculated to within about 5% of its true value by ignoring the volume
changes introduced by the tissue. The volume of CO2 produced can be cal-
culated by multiplying the change in concentration by the entire void vol-
ume of the container.
The significantly lower solubility of oxygen and ethylene in the
aqueous solution of most commodities requires that the volume of the
tissue be subtracted from the container volume when calculating their
production rate.  Flow-Through or Dynamic System

In the flow-through system, tissue is placed in a sealed container through
which a stream of gas flows (Figure 5.4). This flow must be accurately main-
tained during the experiment. After a period of time that is usually equal to
the time required for the flow to replace the container volume three times,
the difference in gas concentrations between the inlet and outlet flows
should be nearly equal to the gases being consumed or produced by the tis-
sue. The system is then in equilibrium, and the differences in concentration
between the inlet and outlet gases will be equal to their production (e.g.,
CO2, C2H4) or consumption (e.g., O2) by the tissue. Gas samples can then
be taken for analysis.
The flow-through system is well suited for long-term experiments and
controlled atmosphere experiments when atmospheres of various composi-
tions are used to ventilate the container. This system has the added advan-
tages that leaks are not critical as long as the gases in the container are
completely mixed, and the physiological active concentrations of evolved
gases can be kept below their critical levels.
To calculate the rate of respiration in a dynamic system, you need to
know the following: weight of tissue, inlet carbon dioxide concentration,
flow of gas, and outlet carbon dioxide concentration.


Gas in


O2 CO2

Carbon dioxide %


0 2 4 6 8 10
(a) (b) Time
Gas out

Figure 5.4  Representation of the setup for measuring O2 consumption or

CO2 production in a dynamic or flow-through system. (a) Depiction of a com-
modity in a flow-through system. (b) Graph showing the increase in CO2 that
would occur as the commodity respired in flow of gas. Note how the CO2
concentration increases to a steady-state level as the loss of CO2 from the
container approaches the production of CO2 by the commodity. Judicious
selection of commodity weight and flow should keep the equilibrium CO2
concentration below around 0.2%.

The production of carbon dioxide or ethylene or the consumption of

oxygen is calculated by multiplying the increase (or decrease) in concen-
tration between the inlet and outlet tubes by the flow rate. Dividing by the
fresh weight (in kg) of the tissue gives the rate of carbon dioxide production
in ml/kg-h.
For example, assume that a 280 gram apple is enclosed in an 843 ml
jar. The carbon dioxide concentration increases from 0.03% to 0.23% as air
flows through the jar at 100 ml/min. It usually takes three volume changes
for a flow-through system to come into equilibrium, so it will take 25.3 min-
utes [(3 × 843 ml)/100 ml/min] for this system to come into equilibrium.
The concentration of carbon dioxide increased 0.20%, so the amount of car-
bon dioxide produced per hour is 12.0 ml (i.e., 0.2% × 100 ml/min × 60
min/h). Dividing by the 0.28 kg fresh weight gives a rate of production of
42.8 ml CO2/kg-h [i.e., (12.0 ml CO2/h)/0.28 kg].
The accuracy of the dynamic method is dependent on knowing the
exact flow rate. The container volume and flow rate are both important in
determining the time it will take for the system to come to equilibrium, but

Respir ato ry Me ta bo lism

as long as the system is at equilibrium, the volume of the container is not

relevant to the calculations. Small leaks are also not important as long as
the gases are thoroughly mixed within the container.

5.5  Sampling and Analyzing

Gas samples can be taken with gastight plastic syringes by inserting the
syringe needle through a septum in the container walls or into flexible tub-
ing protruding from the container. The material used to make most of the
glass and plastic sampling syringes is very impermeable to carbon dioxide
and oxygen, so samples can be held in these containers for days without
significant contamination by gases in the surrounding atmosphere. The
samples are injected in an analyzer and the output compared to standards
to arrive at the correct gas concentration in the sample.
Qualitative and quantitative measures of the component gases in the
sample can be done with a variety of methods and instruments. Early meth-
ods of gas analysis relied on changes in the volume of the gas sample when
specific gases were absorbed. These methods are only of historical interest
today. However, some of the early colorimetric methods are still used. The
reaction of a specific gas with a reagent immobilized on a solid support
causes a color change that identifies the gas and can be used in certain
configurations to estimate the concentration of the gas.
Modern methods of gas analyses rely on electronic instruments.
Some instruments are designed to measure a specific gas. For example, the
infrared analyzer is specific for carbon dioxide, and the paramagnetic or
electrochemical analyzer is specific for oxygen. These instruments do not
require the separation of a gas sample into its individual gas components.
The gas chromatograph separates a gas sample into its individual com-
ponents by passing the sample down a long tube filled with a coated solid
support. The sample is injected into a stream of inert gas (usually nitrogen)
that carries the sample down the tube. The coating and support are selected
to achieve the desired separation of the gases. Upon emerging from the tube,
the separated gases pass through a detector that measures some property
of the gas. Thermal conductivity detectors measure the capacity of the ­sample
to remove energy from a heated filament. Ionization detectors measure the
electrical conductivity of the gas after it has been ionized by ­combustion
(flame ionization detector) or irradiation (photoionization detector).
Samples need not be drawn if the respired gases can be collected. In
a dynamic—or flow-through—system, the commodity is placed in a sealed
container through which a stream of gas is passed. The exit stream is passed
through a column containing caustic solutions or granules that absorb the
respired carbon dioxide. Respiration is determined by subsequent titrimetric


or gravimetric analysis of the material in the column. This system is r­ elatively

easy to set up and is independent of airflow rates, but any leaks in the tubes
leading from the respiration container (which is usually impossible to seal
completely) cause erroneous results. This method is rarely used because
large amounts of plant tissue and long sampling times are required, and the
method is not as accurate as the instrumental methods described previously.

5.6  Instruments and Techniques

Gas chromatograph: All respiratory gases and ethylene can be
assayed with a gas chromatograph by selecting the proper column
and detector. The instrument can be set up to use small (< 1 ml)
samples and rapidly (< 1 min) assay them.
Infrared CO2 analyzer: An instrument that specifically mea-
sures CO2. It can be modified to analyze small (< 1 ml) samples
within 10 s.
Paramagnetic O2 analyzer: An instrument that specifically mea-
sures oxygen. It requires a large sample of gas to analyze and has a
slower response than the other two instruments. Oxygen can also
be measured by polarography or electrochemical means.
Kitagawa tubes: These are glass tubes filled with a specific for-
mulation of reagent to detect a specific gas. A known volume of gas
(e.g., 100 ml) is drawn through the tube, and the reaction between
the reagents and the specific gas component produces a colored
reaction. The length of the colored band is proportional to the con-
centration of the gas in the sample.
Pressure or volume changes: If CO2 produced by respiration
is absorbed by a chemical (e.g., KOH) in a closed container, the
reduction in the volume of O2 will be measurable as a decrease in
pressure. A monometer can indicate the change in pressure by a
small change in volume of a liquid in a U-tube. A large gas sample
(e.g., 100 ml) is usually required, and minor temperature changes
can significantly change the pressure in a sealed container.

5.7  Pre- and Postharvest Factors

Affecting Respiration

5.7.1  Temperature Effects

Without a doubt, the most important factor affecting the rate of respiration
is temperature. Over the physiological range (i.e., 0°C−30°C), increased
temperatures cause an exponential rise in respiration; in general, the

Respir ato ry Me ta bo lism

velocity of a biological reaction increases two- to three-fold for every 10°C

rise in temperature.
The temperature coefficient for a 10°C interval is called the Q10,
which can be calculated by measuring the rate of the respiration (e.g., CO2
production) at two temperatures. Obviously, if the temperature difference
is 10°C, then the Q10 will simply be the quotient of the two rates, that is,
Q10 = R2/R1.
The temperature coefficient is a useful concept, because it allows
calculation of expected respiration rates at one temperature from a known
rate at another temperature using the equation given above. However, the
respiration rate in perishable commodities does not follow ideal behavior,
and the Q10 can vary considerably. At higher temperatures, the Q10 is usu-
ally smaller than at lower temperatures. Typical figures for Q10 are given in
Table 5.1.
These typical Q10 values allow us to construct a table showing the
effect of different temperatures on the rates of respiration or deterioration
and relative shelf life of a typical perishable commodity (Table 5.2).
Table 5.2 shows that if a commodity has a mean shelf life of 13 days at
20°C, it may be stored for as long as 100 days at 0°C and will last no more
than 4 days at 40°C.
Although respiration is normally reduced at low, but nonfreezing tem-
peratures, certain commodities, chiefly those originating in the tropics and
subtropics, show abnormal respiration when the temperature falls below
10°C−12°C. Typically, the Q10 is much higher at these low temperatures

Table 5.1  Typical Q10 Values

Temperature Range (°C) Q10
  0 – 10 2.5 – 4.0
10 – 20 2.0 – 2.5
20 – 30 1.5 – 2.0
30 – 40 1.0 – 1.5

Table 5.2  Effect of Temperature on Rate of Deterioration

Assumed Velocity of Relative Shelf
Temperature (°C) Q10 Deterioration Life
 0  1.0 100
10  3.0 33
20  7.5 13
30 15.0  7
40 22.5  4


than predicted; that is, the rate of respiration falls abnormally rapidly. In
some cases, the respiration rate may even increase dramatically at lower
temperatures, possibly associated with increased glycolysis to compensate
for loss of mitochondrial oxidation. These events are symptoms of the onset
of chilling injury, an economically important low-temperature phenomenon.
Another important respiratory effect of exposure of sensitive com-
modities to chilling temperatures is abnormally high respiration rate (sus-
tained if the tissue has been irreversibly damaged) when the commodity is
returned to normal temperatures. This enhanced respiration presumably
reflects the cells’ efforts to detoxify metabolic intermediates that may have
accumulated during the chilling exposure, as well as to repair damage to
membranes and other subcellular structures.
As the temperature rises beyond the physiological range, the rate
of increase in respiration falls. It becomes negative as the tissue nears its
thermal death point, when metabolism is disorderly and enzyme proteins
are denatured. Many tissues can tolerate high temperatures for short times
(e.g., minutes), and this property is used to advantage in killing some surface
fungi in stone fruits and papaya. Continued exposure to high temperatures
causes phytotoxic symptoms and then tissue collapse. However, condition-
ing and heat shocks (i.e., short exposures to potentially injurious tempera-
tures) can modify the tissue’s responses to subsequent harmful stresses.

5.7.2  Respiratory Quotient

When carbohydrates are the primary substrate respired to produce ATP,
there is an equimolar production of CO2 to O2 consumed. The proportion of
CO2 produced to O2 consumed will vary when other substrates are respired.
Because organic acids (e.g., citric acid) are the product of partially oxidized
glucose, tissue using organic acids in respiration will produce more CO2
per O2 consumed. The ratio of CO2 produced to O2 consumed is called the
­respiratory quotient (RQ) and is defined as the ratio of CO2 evolved to O2 con-
sumed (measured in moles or volumes). RQ values range from 0.7 to 1.3 for
aerobic respiration: carbohydrates have an RQ of around 1.0, while lipids have
an RQ of less than 1.0, and organic acids have an RQ of greater than 1.0. In
the aerobic range, calculation of the RQ can indicate which substrate is being
oxidized during respiration. Very high RQ values can indicate that anaerobic
respiration may be occurring in some parts of the tissues under examination.

5.7.3  Physical Stress

Physical stress has a pronounced effect on the respiration of perishable
products. Both biotic and abiotic stresses cause the tissue to produce a

Respir ato ry Me ta bo lism

signal that migrates from the site of injury and induces a wide range of
physiological changes in adjacent, nonwounded tissue. Some of the more
important changes include enhanced respiration, ethylene production,
phenolic metabolism, and wound healing. Most wound responses are det-
rimental to quality (e.g., rapid softening, browning, toughening), but some
(e.g., wound healing in potatoes) are necessary for their long-term storage.

5.7.4  Internal Factors

A number of commodity parameters are directly related to the respiration
rate and respiratory pattern. Genotype
Some fruits are more perishable than others. You would not, for example,
expect a strawberry to last as long as apples in your fruit bowl. This relative
perishability is usually reflected in the respiration of the different commodi-
ties. The respiration of strawberries is much higher than that of apples.
Some short-lived apple and melon fruit are from cultivars with higher rates
of respiration than long-lived cultivars.  Type of Plant Part

There is a very large variation in the respiration rate among different plant
organs. Storage tissues such as nuts and tubers have low respiration rates.
Tissues with vegetative or floral meristems, such as asparagus and broc-
coli, have very high respiration rates.
As plant organs mature, their rate of respiration typically declines.
This means that commodities harvested during active growth, such as
many vegetables and immature fruits, have high respiration rates. Mature
fruits, dormant buds, and storage organs have relatively low respiration
rates. After harvest, the respiration rate typically declines, slowly in non-
climacteric fruits and storage organs, and rapidly in vegetative tissues and
immature fruits. The rapid decline presumably reflects depletion of respi-
rable substrates that are typically low in such tissues.  Respiratory Climacteric

An important exception to this general decline in postharvest respiration
is the rapid and sometimes dramatic rise in respiration during the ripen-
ing of climacteric fruit. This rise, which has been the subject of intense
study for many years, is normally considered as having four distinct phases:
(1)  preclimacteric minimum, (2) climacteric rise, (3) climacteric peak,
and (4) postclimacteric decline. The division of fruits into climacteric and


nonclimacteric types has been very useful for postharvest physiologists.

However, some fruits, for example, kiwifruit and cucumber, appear to blur
the distinction between the groups. Respiratory rises also occur during
stress and other developmental stages, but a true climacteric only occurs
coincident with ripening.

5.8  Conclusions and Future Perspectives

Respiratory metabolism during postharvest handling and storage should
be carefully regulated to proceed at the minimal rate that maintains prod-
uct quality. While elevated metabolic rates are sometimes needed for short
periods of time to increase quality (e.g., ripening of climacteric fruit, ­curing
of root vegetables), the subsequent maintenance of product quality is opti-
mized at far lower metabolic rates. Storing, transporting, and marketing
harvested horticultural commodities at their lowest tolerated temperature
is the primary method used to suppress metabolism and maximize quality
retention. Controlled and modified atmospheres are used to extend stor-
age life by reducing the oxygen concentration and increasing the carbon
dioxide concentration within the commodities. Postharvest practices could
be formulated that increase the  tissue’s tolerance to lower oxygen and
higher carbon dioxide levels. The significant rise in respiratory metabolism
that accompanies the ripening of climacteric fruit may be physiologically
­unnecessary. Elimination of the respiratory climacteric by either genetic
manipulation or postharvest treatments would significantly increase the
retention of ­storage reserves that could possibly contribute to quality

Kader, A.A. 1992. Postharvest Technology of Horticultural Crops. 2nd ed.
Division of Agriculture and Natural Resources, University of California,
Kader, A.A., and Saltveit, M.E. 2003. Respiration and gas exchange. In
Postharvest Physiology and Pathology of Vegetables, ed. J.A. Bartz and J.K.
Brecht. Marcel Dekker, New York, pp. 7–30.
Kays, S.J., and Paull, R.E. 2004. Postharvest Biology. Exon Press, Athens, GA.
Wills, R.B.H., McGlasson, B., Graham, D., and Joyce, D. 1998. Postharvest:
An Introduction to the Physiology and Handling of Fruit, Vegetables and
Ornamentals. Wallingford, Oxon, UK: CAB International.

Chapter 6

Stomata and
Postharvest Physiology
Uulke van Meeteren1 and Sasan Aliniaeifard 2
1Wageningen University, Wageningen, the Netherlands
2University of Tehran, Tehran, Iran

Abstract 158
6.1 Introduction 158
6.2 Stomata 159
6.2.1 Role of Stomata in Plants 159
6.2.2 Mechanism of Stomatal Closure and Opening 161
6.2.3 Signal Transduction Pathways in Guard Cells
for Stomatal Closure 164
6.3 Role of Stomata in the Postharvest Phase 166
6.3.1 Stomata in Relation to Vase Life of Cut Flowers 166
6.3.2 Stomata in Relation to Quality of Vegetables 169
6.3.3 Stomata in Relation to Quality of Fruits 172
6.4 Preharvest Conditions Leading to Postharvest
Problems via Stomata 174
6.4.1 Relative Humidity and Stomata Control 175 How Does Preharvest Low VPD Affect
Postharvest Stomata Control? 176 Induction of Stomata Morphological
Changes by Low VPD 179


6.4.2 Temperature 181

6.4.3 Light 182
6.5 Stomata and Tolerance to Postharvest Diseases
and Physiological Disorders 184
6.6 Postharvest Treatments and Stomata 186
6.7 Ethylene, Stomata, and Senescence 187
6.8 Conclusions and Future Perspectives 190
References 191

Stomata are pores in the gastight waxy cuticula that covers the outer surface
of aerial parts of plants. They make uptake of CO2 possible, which is needed for
photosynthesis. At the same time, water vapor will leave the plant via the sto-
mata. To optimize photosynthesis, while at the same time preventing excess
water loss, stomata control their opening by a signaling network of pathways
that respond to environmental conditions such as light and darkness, water
vapor pressure deficit (VPD), temperature, CO2, and ethylene. Water loss is
one of the most obvious changes in harvested vegetables, often limiting mar-
keting life, and a negative water balance (uptake of water is insufficient to
compensate transpiration) is one of the most important reasons for the end of
the vase life of cut flowers. Also, the postharvest quality of some fruits is nega-
tively affected by water loss. In leafy vegetables, stomata are portals that make
invasion of bacteria into the inner tissue possible and protect in that way bac-
teria for sanitizers in washing solutions with the risk of food-borne bacterial
diseases. This chapter discusses how several environmental factors, during
preharvest cultivation as well as postharvest storage, influence stomata clos-
ing control in harvested cut flowers, vegetables, and fruits. Also, the role of the
number of stomata and their variability between genotypes and due to cultiva-
tion conditions are discussed in relation to postharvest life. One of the most
striking factors is low VPD (high humidity) during the growth of plants: after
exposure of several days to low VPD, the control of stomata closure is largely
disturbed; stomata do not respond anymore to stimuli that normally induce
closure. This malfunctioning is very persistent and results in high water loss
afterward in the harvested products. Fast cooling of produce can close the
stomata of some crops, while in others, the stomata stay open until wilting.

6.1 Introduction
A waxy cuticle covers the outer surface of the epidermal cells of the a­ erial
parts of plants. This cuticular layer protects plants from drying via its
airtight properties. However, a plant needs gas exchange (intake of CO2

Stom ata a n d P ostharvest P hy s i o l o g y

and O2) for photosynthesis and respiration. Therefore, there are pores in the
cuticle, called stomata, of which the opening can be controlled by surround-
ing guard cells. Stomata are the main openings connecting the internal
leaf space to the outside environment. The intercellular air spaces in plant
tissues are saturated with water vapor, which will exit the tissue through
these stomata. Water loss and respiration are two of the most important
processes involved in postharvest physiology of freshly harvested products;
therefore, understanding the control of the opening of the stomata in the
postharvest phase is of importance to control the postharvest behavior of
plant produce.
Besides gas exchange, stomata also offer opportunities for fungi
and bacteria to enter plant tissues. The infection can take place during the
preharvest growth period as a latent infection that results in postharvest
losses, but contamination with bacteria can also take place during handling
after harvest. Especially when human pathogenic bacteria enter plant tis-
sue via the stomata, they are a serious threat to human health.
The closing and opening behavior of stomata is instantly affected by
many environmental cues. Above this, exposure for some time to specific
environmental conditions can influence the stomatal closing control after-
wards. As a consequence, climatic conditions during cultivation can affect
the stomatal closing control during the postharvest phase. In this chapter,
we describe the control mechanism of stomata control, how this control can
be affected by previous environmental conditions, and the specific possible
consequences for quality changes during the postharvest phase of orna-
mentals, vegetables, and fruits.

6.2 Stomata

6.2.1  Role of Stomata in Plants

The presence of a waxy layer (cuticle) that covers the outer surface of the
epidermal cells greatly reduces water loss from the epidermal cell walls
to the surrounding atmosphere. It also blocks exchange of other gases
between the plant tissue and the air around the plant. Stomata are pores in
the plant epidermis surrounded by a pair of kidney-shaped cells in dicots
(and some monocots) or dumbbell-shaped cells in monocots, the guard
cells (Sack, 1987). These stomata provide an entry channel for CO2 and
an exit for water vapor; this last process is called transpiration. In dicots,
the pattern of stomatal arrangement over the leaf surface is irregular,
while their arrangements are in parallel rows in monocots (Figure 6.1).
The guard cells regulate the opening and closing of stomata to control gas
exchange between a plant and the surrounding environment. The main role




Figure 6.1  Distribution of stomata over the leaf of Tradescantia virginiana

as an example of monocots (a) and Vicia faba as an example of dicots (b).
(Pictures are from author’s archive.)

of stomata is providing enough CO2 for photosynthesis, while at the same

time protecting the water status of the plant by preventing excess water loss
via its opening. They have the capacity of adaptation to the environmental
conditions to minimize water loss while promoting the acquisition of CO2.
Therefore, stomata and their opening and closing control are of vital impor-
tance for the survival of plants. Because of this vital importance, evolution
has resulted in a complex signaling network of pathways that will trigger
opening or closing under certain environmental conditions. This signaling
system comprises positively as well as negatively (inhibiting) interacting
modules (Sirichandra et al., 2009).
Besides photosynthesis and water loss, the temperature of a plant
plays a vital role in plant functioning and survival. High leaf temperatures
can decrease photosynthesis, cause damage to the components of the pho-
tosynthetic apparatus (Schreiber and Berry, 1977; Wise et al., 2004; Camejo
et al., 2005), or ultimately result in the death of tissue. Leaf temperature
depends mainly on the energy going into the leaf (absorbed solar irradia-
tion, absorbed infrared irradiation from surroundings) and energy going
out of the leaf (emitted infrared radiation, heat convection, heat conduc-
tion, heat loss accompanying water evaporation). Because of the role of this
last process, leaf temperature is affected by relative air humidity (RH), leaf

Stom ata a n d P ostharvest P hy s i o l o g y

boundary layer conductance, stomatal conductance, and leaf morphologi-

cal traits, such as the presence of trichomes on the leaf (Jones, 1999; Nobel,
1999). Energy is released when water evaporates at the leaf cell walls to
the intercellular spaces. Energy is required to break down the hydrogen
bonds in the liquid phase of water molecules. This energy is taken from the
leaf and transferred to the water molecules and released as gas molecules.
In the substomatal cavity, the water vapor pressure is in equilibrium with
the apoplast fluid facing the gas phase in the cavity. When water vaporizes
from the substomatal cavity, a new equilibrium establishes between the
water vapor pressure of the cavity and cells. Consequently, the water vapor
and associated energy are released into the atmosphere via the stomata
(thermal dissipation). Therefore, transpiration of water vapor from the sto-
mata is associated with cooling of the leaf (Hetherington and Woodward,
2003). For example, using infrared thermal imaging, it has been shown that
mutants with open stomata have lower leaf surface temperature than wild-
type plants (Merlot et al., 2002). Therefore, proper functioning of stomata
is vital for balancing leaf temperature and water loss. In Arabidopsis plants
that had developed at high temperature (28 ° C), increased water loss was
associated with an enhanced leaf cooling capacity. However, the leaves of
the high-temperature-developed plants possess lower stomatal density and
reduced stomatal size. In this case, to cool down the leaf, plant architec-
tural adaptions such as petiole elongation, leaf elevation above the soil, and
decrease in leaf thickness enhanced the diffusion of water vapor from the
stomata (Crawford et al., 2012; Murata and Mori, 2013).
As another role of stomata, it is generally accepted that the tran-
spirational flux is the main mechanism of transport of nutrients in plants
(Mengel and Kirkby, 1982; Novák and Vidovič, 2003).

6.2.2  Mechanism of Stomatal Closure and Opening

Closure and opening of the stomata pores depend on many factors, including
environmental factors such as light, temperature, and RH (or better water
VPD), CO2 concentration inside the stomatal cavity, water availability, and
pathogens, and endogenous factors such as phytohormones and their inter-
actions, and secondary messengers. Stomatal aperture changes over diur-
nal cycles. To facilitate CO2 assimilation, stomata stay open during the day,
especially in response to blue light, and tend to be closed at night (Talbott
and Zeiger, 1998; Schroeder et al., 2001; Tallman, 2004). However, to con-
serve water, plants with a crassulacean acid metabolism (CAM) close their
stomata during the daytime and open at night to uptake CO2 (Bohnert et al.,
1995; Black and Osmond, 2005). In this way, CAM plants lose less water and
are adapted to dry conditions. The number of stomata over the leaf surface
is species dependent (Table 6.1) and highly depends on the environmental


Table 6.1  Stomatal Density on the Leaf of Some Horticultural Crops

Stomatal Density
Species (no. mm−2) Reference
Faba bean 30–50 Aliniaeifard et al. (2014)
Rose 37–65 Torre et al. (2003), Fanourakis et al.
(2011, 2013a), Giday et al. (2013a)
Tradescantia 19–22 Rezaei Nejad and van Meeteren (2005)
Cucumber 322–460 Bakker (1991), Savvides et al. (2012)
Tomato 75–203 Willmer and Johnston (1976), Bakker
(1991), Amjad et al. (2014)
Eggplant 136–182 Bakker (1991)
Arabidopsis 100–600 Doheny-Adams et al. (2012)
Anthurium 100–220 Schroeder and Stimart (2005)
Strawberry 120–330 Blanke (2002), Orsini et al. (2012)
Papaya 265–400 Parés et al. (2008)
Basil 50–250 Barbieri et al. (2012)
Almond 150–326 Camposeo et al. (2011)
Potato 80–260 Yan et al. (2012)
Gerbera 116–194 Romero-Aranda et al. (1994)
Azalea 108–163 Ceulemans et al. (1984)
Dodonaea 320–430 Shtein et al. (2011)
Apple 320–575 Blanke (1987), Blanke and Lenz (1989),
Blanke et al. (1994)
Trigonella 225–245 Willmer and Johnston (1976)
Orange 360–500 Moreshet and Green (1980), Burg
Banana 1700 Wardlaw (1936)

condition prevalent during growth of the plants. The distribution of stomata

over the leaf surface is not a random process. They are distributed in a way
that effective diffusion of the gases between leaf internal tissues and the
surrounding environment is provided. Therefore, development of stomata
must be conserved in a good order to support their roles. According to the
one-cell spacing rule (Sachs, 1991), around the ­stomata is always at least
one nonstomatal epidermal cell that separates stomata from each other.
Since stomatal opening and closure require efficient trafficking of water
and ion molecules between guard cells and neighboring nonstomatal cells,
the one-cell spacing rule is vital for proper regulation of stomatal aperture
(Peterson et al., 2010). Stomatal opening and closing are controlled by guard

Stom ata a n d P ostharvest P hy s i o l o g y

cell swelling and shrinking, respectively. Guard cell turgor dynamically

changes in response to environmental conditions and endogenous signals.
Since guard cells lack plasmodesmata, most of the effluxes and influxes of
the solutes are accomplished through pumps, ion channels, and transport-
ers located in the plasma membrane (Daszkowska-Golec and Szarejko,
2013). Stomatal opening is initiated by extrusion of protons from guard cell
membranes through H+ -ATPases. Signals such as blue light and auxin are
positive regulators of H+ -ATPases, while calcium and the phytohormone
abscisic acid (ABA) operate as negative regulators of H+ -ATPases (Hentzen
et al., 1996; Parvathi and Raghavendra, 1997; Elzenga and Van Volkenburgh,
1997; Frechilla et al., 1999; Leonhardt et al., 1999; Elzenga et al., 2000; Zeiger,
2000; Zhang et al., 2004, 2007; Takemiya et al., 2006; Marten et al., 2007a;
Merlot et al., 2007; Su et al., 2007; Harada and Shimazaki, 2009; Zhao et al.,
2012). Proton extrusion induces plasma membrane hyperpolarization and
apoplast acidification. The created voltage gradient activates inward-recti-
fying K+ channels. Influx of K+, which is balanced by the influx of counter-
ions such as Cl− and NO3− and inorganic solutes such as malate, enhances
guard cells’ osmotic potential. Therefore, water pumped into the guard cells
causes swelling of the guard cells, and as a result, stomatal opening occurs.
Stomatal closing is initiated by inhibition of the H+ -ATPases, which depolar-
izes the plasma membrane. Following depolarization, outwardly rectifying
K+ channels enhance the driving force for K+  efflux and decrease the K+
level inside the guard cells. Two distinct types of depolarization-activated
anion channels act in the plasma membrane of the guard cells during sto-
matal closure: one activates rapidly by depolarization and also deactivates
rapidly by hyperpolarization (R-type anion channels), and the other one
activates and deactivates very slowly by voltage gradients (S-type anion
channels) (Schroeder and Keller, 1992). Anion channels facilitate efflux
of malate2−, Cl−, and NO3− from the guard cells. Moreover, gluconeogenic
conversion of malate2− into starch would result in further decrease of the
malate2− level. Cytosolic Ca 2+ elevation (also oscillation) through channels
located in the plasma membrane and in the tonoplast usually accompanies
stomatal closure (Leckie et al., 1998; Siegel et al., 2009; Hubbard et al., 2012;
Daszkowska-Golec and Szarejko, 2013).
Efflux of K + and Cl− ions or malate2− through guard cells’ membrane
decreases osmotic potential, and as a result, water exits from the guard
cells, which causes them to shrink, resulting in stomatal closure (Blatt,
2000; Schroeder et al., 2001; Outlaw, 2003).
Plants dynamically respond to changes in environmental conditions
by regulating the aperture of the stomata. Plant responses to environmen-
tal stresses (e.g., drought) are usually associated with induction of abscisic
acid (ABA) production. ABA, through its signal transduction pathway,
causes stomatal closure (Hu et al., 2006; Endo et al., 2008; Lee and Luan,
2012; Sreenivasulu et al., 2012).


6.2.3  Signal Transduction Pathways

in Guard Cells for Stomatal Closure
Guard cells perceive multiple signals from the environment and integrate
them to internal signals. By following complex transduction pathways, guard
cells respond by regulating stomatal aperture in order to adapt to the envi-
ronment (Bohnert et al., 1995; Qin and Zeevaart, 2002; Lebaudy et al., 2008;
Oh et al., 2009; Kim et al., 2010; Trontin et al., 2011; Lee and Luan, 2012; Zhu
et al., 2012; Christmann et al., 2013; Kuromori et al., 2014). Over the past
several years, many internal signals have been recognized in guard cells in
response to different environmental signals. For example, calcium, reactive
oxygen species (ROS), phosphatidic acid, cyclic guanosine 3',5’-monophos-
phate (cGMP), nitric oxide (NO), and pH have been found as essential sig-
nals mediated in stomatal closure (Suhita et al., 2004; Li et al., 2006; Wang
and Song, 2008; Xue et al., 2009; Dubovskaya et al., 2011; Kim et al., 2010;
Stael et al., 2011). Although an ABA-independent pathway for closure of the
stomata has also been proposed (Yoshida et al., 2006; Huang et al., 2009;
Montillet et al., 2013; Roychoudhury et al., 2013), ABA is considered to be the
main phytohormone that promotes stomatal closure, which helps to mini-
mize water loss by decreasing transpiration via stomata. This function of
ABA is accomplished through modulating a complex and sophisticated cas-
cade of biochemical and molecular events (Hauser et al., 2011). Almost all of
the previously mentioned internal signals are involved in guard cells’ ABA
signaling pathway for closure of the stomata (Leung et al., 1997; Kwak et al.,
2002; Suhita et al., 2004; Li et al., 2006; Zhu et al., 2007; Neill et al., 2008; Wang
and Song, 2008; Hubbard et al., 2010, 2012; Kim et al., 2010; Dubovskaya
et al., 2011; Joshi-Saha et al., 2011; Hossain et al., 2011). The ABA-induced
stomatal closure is often associated with an increase in guard cells’ calcium
concentration. However, calcium-dependent and ­calcium-independent ABA
signaling pathways have been suggested for ABA-induced stomatal closure
(Figure 6.2) (MacRobbie, 1990; Li and Assmann, 1996; Levchenko et al.,
2005; Sutter et al., 2007; Marten et al., 2007b; Geiger et al., 2009; Siegel et al.,
2009; Geiger et al., 2010; Joshi-Saha et al., 2011).
To initiate ABA signal transduction, guard cells are equipped with ABA
receptors to bind to ABA (Moes et al., 2008; Fujita et al., 2009; Ma et  al.,
2009; Santiago et al., 2009; Cutler et al., 2010; Raghavendra et al., 2010; Lee
et al., 2013). Binding of ABA with its receptors inhibits the activity of group
A protein phosphatases 2C (PP2Cs) (Moes et al., 2008; Fujita et al., 2009; Ma
et al., 2009; Park et al., 2009; Santiago et al., 2009; Raghavendra et al., 2010).
OST1 is an Arabidopsis SnRK2-type protein kinase that, together
with several other SnRK2-type protein kinases, is also known to func-
tion in ABA responses (Belin et al., 2006; Fujita et al., 2009; Lee et al.,
2013; Yoshida et al., 2002, 2006). In contrast to A-type PP2Cs, SnRK2-type

Stom ata a n d P ostharvest P hy s i o l o g y

protein kinases are positive regulators of ABA signaling (Fujii et al., 2009;
Fujii and Zhu, 2009; Ma et al., 2009; Park et al., 2009; Umezawa et al., 2009;
Vlad et  al., 2009; Lee and Luan, 2012). Downstream of ABA receptors,
PP2Cs, and SnRKs are ion channels that control stomatal movements (Fujii
et al., 2009; Geiger et al., 2009; Lee et al., 2009). The guard cell slow-type
anion channel (SLAC1) may represent an essential component for stomatal
closure induced by ABA or other signals (Negi et al., 2008; Vahisalu et al.,


ABA Receptors
Calcium-independent ABA signaling

Calcium-dependent ABA signaling



Ion channels
(e.g., SLAC1)


Figure 6.2  Simplified ABA signal transduction pathway in guard cells for
closure of the stomata. In a calcium-independent ABA signaling pathway,
perception of ABA by receptors leads to inactivation of type 2C protein
phosphatases (PP2C). As a result, S-type anion channels (SLAC1) will be
activated by a SnRK2-type protein kinase (OST1). Consequently, stomatal
closure occurs. In the calcium-dependent ABA signaling pathway, calcium-
dependent protein kinases (CDPKs) can induce stomatal closure via activa-
tion of SLAC1 (Data from Hubbard, K.E. et al., Genes and Development
24, 1695–1708, 2010; Kim, T.H. et al., Annual Review of Plant Biology
61, 561–591, 2010; Antoni, R. et al., Current Opinion in Plant Biology 14,
547–553, 2011; Sreenivasulu, N. et al., Gene 506, 265–273, 2012; Lee,
S.C. et al., Molecular Plant 6, 528–538, 2013).


2008, 2010; Geiger et al., 2009). SLAC1 acts as a substrate for and is acti-
vated by OST1 (Geiger et al., 2009; Lee et al., 2009, 2013).
Calcium-dependent protein kinases (CDPKs) function as essential
elements of the calcium-dependent ABA signaling (Zhu et al., 2007). It has
been shown that SLAC1 can be activated by CDPKs, which leads to stoma-
tal closure (Mori et al., 2006; Geiger et al., 2010) (Figure 6.2).

6.3  Role of Stomata in the Postharvest Phase

As stomata play an important role in controlling gas exchange between
internal plant tissues and the surrounding environment, they can affect the
postharvest physiology of plants and plant products as far as postharvest
physiology is influenced by gas exchange. In this chapter, we discuss spe-
cific postharvest issues of cut flowers, vegetables, and fruits that are related
to the behavior of stomata in these products.

6.3.1  Stomata in Relation to Vase Life of Cut Flowers

The vase life of cut flowers ends when the flowers have lost their ornamen-
tal value. Most flowers attached to the plant lose their ornamental value
because of senescence symptoms as a result of their advanced development;
they show changes in color, wilting of petals, closing of flowers, or abscis-
sion of flowers or petals. However, when flower shoots are cut and placed
in water, they often lose their attractiveness for the consumer because of
premature wilting of flowers or leaves, or loss of stem turgidity; these are
all symptoms of a water deficit (Van Doorn, 1997, 2012). This water deficit
is a result of a negative water balance: the rate of water uptake has become
less than the rate of water loss by transpiration. Other consequences of the
negative water balance for the quality of cut flowers can be impairment of
flower opening and precocious senescence.
In most cases, the imbalance between uptake and loss of water is
the result of a hampered water uptake, due to a gradual decrease in the
hydraulic conductance of the cut flower stem after cutting (Aarts, 1957; van
Meeteren, 1978, 1989; Rogers et al., 1979; Halevy and Mayak, 1981). There
can be several reasons for this decrease in stem hydraulic conductance:
growth of bacteria at the cut stem surface or inside the xylem vessels,
uptake of air into the cut-open xylem vessels at the cut stem surface (air
embolism), wound-induced responses in the cut stem, or formation of vapor
cavities in the xylem fluid (cavitations) (Tyree and Dixon, 1986; Williamson
and Milburn, 1995; Van Doorn, 1997). Because of the ongoing transpira-
tion of the cut flower, the decrease in hydraulic conductance of the stem
results in a decrease of the water potential of the flower and loss of turgidity.

Stom ata a n d P ostharvest P hy s i o l o g y

The decrease in water potential can cause cavitations, which will further
enlarge the imbalance between water uptake and loss.
The negative effect of a decreased stem hydraulic conductance on the
water balance of a cut flower will be postponed or less severe when the tran-
spiration rate of the cut flower is low. As an example, it has been shown that
the vase life of cut roses can be prolonged by various ways that lower their
transpiration: cooling, high air humidity, and leaf removal (Zieslin, 1989).
Mayak and Halevy (1974) showed that the difference in vase life between a
short-lived cultivar and two long-lived cultivars of rose was large, especially
under conditions promoting high transpiration rates, and was narrowed when
either flowers were held in mild conditions or the leaves were stripped off.
Most cut flowers include, besides one or more flowers, a stem that
often bears leaves. The main parts of a flower consist of a vegetative part,
containing sepals and petals, and a reproductive part, containing stamens
and a gynoecium. Because a waxy cuticle covers the outer surface of the
epidermal cells of leaves, sepals, petals, and the gynoecium, most of the
transpiration of cut flowers is via the stomata. Therefore, it can be expected
that the number and the closing and opening behavior of stomata play a role
in vase life. Stomata are usually present in green epidermal tissues such as
leaves, sepals, and green (parts of) petals (Chimona et al., 2012), but also in
some nongreen petals, stomata can be found (Effmert et al., 2005; Chimona
et al., 2012). Stomata can also be completely absent in petals (Mayak and
Halevy, 1974; Tahir and Rajput, 2010). In petals of commercially important
crops like roses (Rosa hybrida), carnations (Dianthus caryophyllus), and ger-
bera (Gerbera jamesonii), stomata were not found (Van Doorn, 1997). As
shown in the overview by Van Doorn (1997), numbers of stomata in petals
vary between 0 and 45 per cm 2, which is very low compared with the num-
bers of stomata found in leaves, which varies between 5 and 1000 per mm 2,
depending on species and growth conditions (Table 6.1) (Hetherington and
Woodward, 2003). When present, stomata in petals are not always func-
tional, as shown for several orchids (Hew et al., 1980). An exceptional flower
in this context is tulip; tulip petals can contain 5–49 stomata per mm 2, and
a temperature-dependent stomatal movement in tulip petals controls water
transpiration during flower opening and closing (Azad et al., 2007). In con-
clusion, for most cut flowers, the main transpiration is via the stomata in the
leaves of the cut flowering stem.
The role of stomata in vase life was elegantly demonstrated by Halevy
et al. (1974) in an experiment about the effects of application of ABA on
the vase life of cut roses. Application of ABA can induce stomatal closure,
reduce water loss, and extend the vase life of cut rose flower shoots bear-
ing leaves. Its effect was very pronounced when the flowers were exposed
to conditions that promote transpiration (28°C and 45% RH). However, in
a stomata-less system (leafless flower shoots) or in leafy shoots held in
darkness when all stomata were closed, ABA shortened vase life because it


increased the aging of the flowers and some biochemical processes associ-
ated with it (RNase activity and reduction in protein content).
Elibox and Umaharan (2008) evaluated the vase life of 26 anthurium
(Anthurium andraeanum Hort.) cultivars. Among the tested cultivars, there
were large differences in vase life, varying from 14 to 49 days. Anthurium cut
flowers do not hold leaves on their cut stem and have a low transpiration rate
and long vase life. Still, symptoms associated with the end of the vase life in
anthurium are typical of water stress (Paull, 1982). Anthurium cut flowers
have a brightly colored modified leaf, the spathe, which subtends a spadix
with more than 300 spirally arranged minute flowers. This spathe contains
stomata, mostly on the abaxial side, of which the density varies between
the cultivars from 1.8 to 25.7 per mm 2. Although the rate of water loss of
anthurium cut flowers is relatively low, of 12 morphophysiological character-
istics, only spathe color and abaxial stomatal density were able to accurately
predict vase life (Elibox and Umaharan, 2008). In their study, Elibox and
Umaharan (2008) also showed that changes in vase life of cultivars over
season were mediated through changes in stomatal density. However, culti-
var differences in stomata density of the spathe explained only a small pro-
portion of cultivar variation in vase life. In a later study, the same authors
showed that the balance between factors affecting water uptake, which is
determined by the timing, extent, and duration of vascular occlusion, and
those affecting water loss (e.g., stomatal density and regulation) may con-
tribute to cut flower senescence in anthurium (Elibox and Umaharan, 2010).
In an evaluation of snapdragon (Antirrhinum majus) cultivars, that
differed in length of vase life, leaf stomatal numbers and postharvest water
loss were shown to be important factors in vase life. Cut flowers of a geno-
type with 9 days longer vase life had 53% fewer leaf stomata (Schroeder
and Stimart, 2005). However, long vase life was especially associated with
an early reduction in transpiration followed by low, steady transpiration,
while short-lived genotypes had a linear transpiration pattern over the vase
period. This indicates that differences in stomatal control (opening and
closing) of water loss between cultivars are likely to be a more important
factor than the number of stomata (Schroeder and Stimart, 2005). This
agrees with a previous paper of Rutland et al. (1987), in which they showed
that stomatal density did not correlate with vase life and transpiration of 10
different snapdragon cultivars.
Number of stomata (stomatal density) is not only dependent on geno-
type (cultivar), but also affected by the environmental conditions during
the cultivation of the cut flowers. Four of five rose cultivars tested showed
a significantly higher number of stomata per cut flower at higher cultiva-
tion temperatures (Pandey et al., 2007). However, the vase life of the cut
flowers was not affected by the preharvest temperature conditions (Pandey
et al., 2010). This strengthens the conclusion that stomata number per se
is not the determining factor for vase life. High air humidity (RH) during

Stom ata a n d P ostharvest P hy s i o l o g y

cultivation greatly reduced the vase life of cut roses, which was accompa-
nied by high transpiration rates (Torre, 1999; Mortensen and Gislerod,
2000) and symptoms that are typically related to water stress, including
premature flower and leaf wilting, as well as pedicel bending (Mortensen
and Gislerod, 2000; Torre and Fjeld, 2001; In et al., 2007). Roses grown at
high RH (90–95%) showed a significantly higher number of stomata and
longer stomata, as well as wider stomatal apertures, than roses grown at
a moderate RH of 60%–70% (Torre et al., 2003; Fanourakis et al., 2013a).
However, in rose leaflets subjected to desiccation, the enhanced stomatal
conductance in high-RH- as opposed to moderate-RH-grown plants was
mostly due to poor stomatal closure and, to a lesser extent, the combined
result of higher stomatal density and longer pore length (Fanourakis et al.,
2013a). This confirms that the reduced degree and, especially, the reduced
rate of stomatal closure are the primary causes of the large genotypic varia-
tion in the control of water loss in high-RH-grown rose plants. It is known
that stomata respond rapidly to air humidity, resulting in higher stomatal
conductance at high RH (Hall et al., 1975; Morison and Gifford, 1983).
Growth of rose plants at high RH, however, resulted in stomata that are
less responsive to desiccation. As a result, transpiration of the cut flowers
remains also high when the water potential in the cut flower decreases. This
implies that cultivation of cut flowers at high RH makes the flowers more
vulnerable to dry storage or transport, and to a hampered water uptake at
the consumer’s, caused by a decreased hydraulic stem conductance of the
cut flower. A limited exposure period to high RH of a few days can already
disturb the stomata closure response to desiccation (Rezaei Nejad and van
Meeteren, 2008; van Meeteren et al., 2009; Aliniaeifard et al., 2014). This
suggests that a period of some days of high RH during the preharvest stage
can have a large impact on the postharvest quality of cut flowers.

6.3.2  Stomata in Relation to Quality of Vegetables

In this chapter, we use the nonbiological definition of vegetables, which is
based on culinary and cultural traditions. In culinary terms, a vegetable is
an edible plant or its part, intended for cooking or eating raw (Vainio and
Bianchini, 2003). In botanical terms, a vegetable can be a root, stem, leaf,
immature flower bud, fruit, or sprout. Fruit vegetables often have a savory
flavor, while culinary fruits (discussed in the next part) often have a sweet
taste and are used for desserts or as a snack. As discussed before, stomata
can be found in most green plant parts. That implies that root-type vegeta-
bles, such as carrot, sweet potato, beet, and radish, are vegetables without
stomata. As the majority of stomata are on the leaves of plants, leafy veg-
etables will have a large number of stomata. But we can also find stomata in
many fruits, especially in green fruits, as most immature fruits are. Fruits of


cucumber, okra, and several legumes, such as pea and bean, are used in an
immature stage as a vegetable. The stomata number on pickle cucumbers
varies from 10 to 100 mm−2 (Smith et al., 1979), what is comparable to some
leaves (see before). The stomatal density decreases when the cucumber
fruit size increases, because stomata are differentiated in an early stage of
fruit development. So, the total number of stomata per fruit does not change
during growth of the fruit. In the legume plants, pots are also rich in sto-
mata. In the pot of broad bean or pea, stomata in the exocarp are 25% of the
stomatal density on the leaf abaxial surface and 50% of the stomatal density
on the adaxial side of the leaf (Willmer and Johnston, 1976; Atkins et al.,
1977; Blanke and Lenz, 1989). The stomatal density in broad bean is around
19 stomata mm−2, and in pea is around 40 stomata mm−2. It has been shown
that 12,000–14,000 stomata exist on one pea pod. The maximum density of
stomata in the pea can be observed after anthesis (Blanke and Lenz, 1989).
Water loss is one of the most obvious changes in harvested ­vegetables,
often being the factor that limits marketing life (Ben-Yehoshua and Rodov,
2003; Shamaila, 2005). Therefore, it is surprising that the effects of prehar-
vest growth conditions and postharvest handling practices on the stoma-
tal aperture of harvested vegetables have received little attention. Water
loss during storage can be controlled with proper and rapid precooling and
storage humidification. In chilling-sensitive vegetables, however, control of
water loss is more difficult, since they can only be stored at warmer tem-
peratures. Also, the environmental conditions in different links of a supply
chain are not always under strict control, which makes the role of stomata
of even more importance. In a study by Thomson (2005), observations
were made on stomatal apertures in the leaves of Japanese mustard spin-
ach (Brassica rapa var. perviridis ‘Komatsuna’) and a large India mustard
(Brassica juncea ‘Red Giant’) in response to cooling and holding treatments
representing commercial procedures. When the leaves were held in air at
20°C, the stomatal aperture of both Brassicas progressively reduced after
harvest; after 15 min, closure was largely complete in B. juncea, while in
B. rapa, stomata closure took about 30 min. Stomata of B. juncea also closed
at 4°C; however, 4°C inhibited stomatal closure of B. rapa. Besides cool-
ing by air, following harvest, B. rapa leaves were also dipped for 20 min in
iced water and in water at 20°C. Neither water dips interfered with stoma-
tal ­closure. However, after these dips and a further 25 min in air at 20°C,
stomata remained largely closed for iced leaves, but had begun to open on
leaves dipped in the 20°C water.
Reducing water loss in B. juncea not only prevented wilting, but
also reduced leaf yellowing; it increased the sweetness and retarded pro-
tein degradation and the loss of vitamin C (Lazan et al., 1987a, 1987b).
Also, in lettuce, the retention of water played an important role in main-
taining ascorbic acid content (Agüero et al., 2011). In general, the nutri-
tional value of vegetables is very much affected by water loss (Paull, 1999;

Stom ata a n d P ostharvest P hy s i o l o g y

Ben-Yehoshua and Rodov, 2003; Shamaila, 2005). However, there has only
been limited research evaluating the impact of water loss on the nutritive
value of vegetables. Ezell and Wilcox (1959, 1962) found that water loss was
associated with significant losses of both vitamin C content and carotene in
a wide range of vegetables (kale, collards, turnip greens, spinach, cabbage,
and snap beans). Diminishing water loss of broccoli by misting resulted in
significant retention of ascorbic acid (Barth et al., 1992). Also, preventing
water loss of strawberries (by wrapping) reduced the loss of ascorbic acid.
As modifications of O2 and CO2 levels were only minimal in the wrapped
treatments, the effect was not due to their modification (Nunes et al., 1998).
Water loss is clearly an important factor in the retention of quality, includ-
ing nutritional quality, in vegetables. As discussed in Section 6.4, the culti-
vation of plants at low VPD can influence the closing ability of stomata. In
that way, the VPD during the preharvest phase of vegetables influences the
postharvest life and nutritional quality of leafy vegetables via the stomata. A
higher rate of water loss after harvest of plant leaves was observed in basil
(Ocimum basilicum L.) and lemon balm (Melissa officinalis L.) as a result
of exposure during cultivation to low VPD than in plants grown at higher
VPDs (Islam et al., 2010). Postharvest life was negatively correlated to the
rate of water loss via stomata after harvest of the plants (Islam et al., 2010).
Whether VPD during the growth of legume plants also affects the stomatal
closing ability of the pots is unknown.
Some recent outbreaks of bacteria-related diseases after consumption
of fresh vegetables have increased the interest in food-borne pathogens and
especially in bacteria–plant interactions. Since stomata are the portals into
the plant for many bacterial species that cause plant diseases, there is more
and more interest in the role that stomata play in food safety, especially
in leafy greens. Scanning electron microscopy studies of lettuce leaves
showed that bacteria can infiltrate stomata and in that way are protected
from chemicals such as sodium hypochlorite during the washing of the
leaves (López-Gálvez et al., 2010). Therefore, washing can be ineffective in
eliminating Escherichia coli cells on leaf surfaces, and such is also the case
for sanitizers used in the washing solution. The main function of sanitizers
in the washing of fresh-cut products is to avoid cross-contamination.
Incubation of GFP-tagged Salmonella enterica with iceberg lettuce
leaves in the light resulted in aggregation of bacteria near open stomata
and invasion into the inner leaf tissue (Kroupitski et al., 2009). In contrast,
when stomata were closed due to darkness, incubation resulted in a scat-
tered attachment pattern and very poor stomatal internalization. However,
Salmonella internalization was not enhanced by forcing stomatal opening
in the dark by the use of fusicoccin. These results imply not only that light
was necessary for stomata opening, but also that the pathogen is attracted
to nutrients produced de novo by photosynthetically active cells. Some plant
pathogens force stomata to open after they have closed as part of the plant’s


initial response to those invasive bacteria. However, lettuce stomata do not

close in response to Salmonella, probably because this bacterium is not a
plant pathogen (Kroupitski et al., 2009). Also, a few listeriosis outbreaks
have been linked to consumption of contaminated vegetables. By dipping
plants of Arabidopsis thaliana for 5 min into water containing Listeria mono-
cytogenes expressing GFP and rinsing repeatedly, it was demonstrated that
these bacteria can internalize the plants via their stomata (Milillo et  al.,
2008). The bacteria were visualized in the intercellular spaces of the leaves
and recovered from the leaves at densities from 1.52 to 2.17 log CFU cm−2 .
Ten days after exposure, bacterial numbers had increased over initial
numbers by 2.60–2.95  log  CFU cm−2 . Taking measures to limit the pen-
etration process of bacteria in leafy vegetables via stomata could improve
food safety.

6.3.3  Stomata in Relation to Quality of Fruits

It is generally accepted that the main plant structure responsible for gas
exchange between a plant and its environment, stomata, is distributed over
the leaf surface. However, some structures similar to the leaf stomata have
been found in the fruits of many plants. Increased fruit transpiration at
light and decreased transpiration at darkness, together with the morphol-
ogy of a fruit’s guard cells, confirm that fruit’s stomata can function like
those on the leaves (Blanke and Lenz, 1989). Fruits of some horticultural
crops possess stomata, while others lack them. The number of stomata
over the fruit surface is species dependent (Table 6.2) and depends highly
on the stage of the fruit development and the environment. For some crops,
different results regarding the existence of stomata have been reported.
For example, the presence of stomata on pepper fruit has been reported
by Fallik et  al. (1999), while Smith et al. (2006) reported no stomata on
pepper fruit. In some fruits, the stomata are not evenly distributed over
the fruit surface. For instance, the stylar scar region of sweet cherry has
the highest stomatal density, followed by the cheek, while the stem cavity
region is lacking stomata (Peschel et al., 2003). Stomata in sweet cherry
are directly or indirectly involved in water transport. When stomata are
functional, their role in water transport varies during the course of a day,
and they may be relevant to the process of fruit cracking (Christensen,
1976; Peschel et al., 2003).
In some crops at early stages of fruit development, stomata exist on
the fruits (especially when they are green), while with progress toward rip-
ening their number decrease or sometimes they disappear at the late stage
of fruit development (Blanke and Lenz, 1989; Rogiers et al., 2004). Grape
berries contain some functional stomata at the preveraison stage, but they
become clogged with wax and nonfunctional in the postveraison stage of

Stom ata a n d P ostharvest P hy s i o l o g y

Table 6.2  Stomatal Density on the Fruits of Some Horticultural Crops

Species Stomatal Density Reference
Strawberry 1 – 6 (no. mm )
−2 Blanke (2002)
Cucumber 20 – 30 (no. mm−2) Adams and Ho (1995)
Tomato –1 to 0 (no. mm−2) Willmer and Johnston (1976),
Adams and Ho (1995)
Grape 8 – 10 (per berry) Blanke and Leyhe (1988)
Sweet cherry 143 – 2,124 (per fruit) Peschel et al. (2003)
Apple 600 – 6,000 (per fruit) Blanke and Lenz (1989); Blanke
1 (no. mm−2) (1994)
Faba bean 19 ± 2 (no. mm−2 on pod) Willmer and Johnston (1976),
Blanke and Lenz (1989)
Pea 40 (no. mm−2 on pod) Blanke and Lenz (1989)
Orange 6 – 75 (no. mm−2) Turrell and Klotz (1940),
Moreshet and Green (1980),
Blanke (1994)
Banana 3.2 – 11 (no. mm−2) Wardlaw (1936), Johnson and
Brun (1966)
Avocado 20,000–30,000 (per fruit) Blanke (1992, 1994)
3 – 5 (no. mm−2)
Trigonella 61 ± 2 (no. mm−2 on pod) Willmer and Johnston (1976)

fruit development (Rogiers et al., 2004). The transformation from functional

stomata at the preveraison stage to nonfunctional stomata (lenticel covered
by wax) at the postveraison stage causes a decrease in berry transpirational
water loss at the late stage of fruit ripening (Rogiers et al., 2004). Therefore,
the flesh of the grape berry at the postveraison stage is highly isolated from
the outside environment, causing higher CO2 and lower O2 concentrations
in the berry core than in fruits with a high number of stomata, such as pear
(Rogiers et al., 2004; Franck et al., 2007; Mencarelli and Bellincontro, 2013).
The berries are attached to the stem through a pedicel. The pedicel is cov-
ered with a thin layer of cuticle and contains a high number of stomata and
lenticels. These properties make the pedicel vulnerable to water loss when
the RH of the surrounding environment is low. High permeability of the
pedicel and its high water loss properties induce a water potential difference
that draws water from the berry, leading to water loss from the airtight ber-
ries (Mencarelli and Bellincontro, 2013).
Young apple fruits are covered by a thin cuticle after petal fall. In this
stage, stomata are completely formed and uniformly distributed over the
fruit (Schwertfeger and Buchloh, 1968). The number of stomata in apple
fruit is determined before petal fall. Stomatal density in apples is reported


between 10 and 20 stomata mm−2 around the petal fall (Blanke, 1986; Blanke
and Lenz, 1989). Then after 1 week, it decreases to 6–8 stomata mm−2 , and
thereafter, when the expansion of the cuticle occurs, it decreases further to
less than 1 mm−2 (Schwertfeger and Buchloh, 1968; Blanke, 1986; Blanke
and Lenz, 1989). It is reported that an apple fruit with 10 mm diameter
would have around 600–6000 stomata per fruit (Table 6.2) (Blanke and
Lenz, 1989).
Also in Valencia oranges, the stomatal density is at its maximum
(160  mm−2) when the fruit is small. Fruit growth expands the space
between the stomata; in large fruits, the number of stomata is decreased
to 20–50 mm−2 (Moreshet and Green, 1980). In navel oranges, the stoma-
tal density (10–16 mm−2) is lower than that in the fruit of Valencia orange
(Turrell and Klotz, 1940). It seems that in all fruits, even when they are ripe,
stomata exist on them, but they are covered by a wax layer.
In banana fruit, contrary to the leaf, the stomata are not distributed in
a parallel way. The number of stomata is between 3 and 11 mm−2 in imma-
ture fruits and 3 and 6 mm−2 in full and mature fruits (Johnson and Brun,
1966). The stomatal density in the leaf (Table 6.1) of banana (1700 mm−2) is
considerably higher than their numbers on the fruits (Table 6.2) (Wardlaw,
1936). Although many studies reported nonfunctional stomata in banana
fruit, it has been shown that just before the harvest, they are functional in
response to light and CO2 and capable of gas exchange with the environ-
ment (Moreshet and Green, 1980; Burg, 2004).
Stomatal density in tomato fruits is very low, and in some studies,
even no stomata were reported on the tomato fruit, while the number of sto-
mata in cucumber is relatively high (Table 6.2). Higher stomatal densities
correlate positively with higher water loss from the fruits. Measuring the
amount of water loss immediately after harvest showed a 20 times higher
water loss for cucumber fruits than for tomato fruits (Adams and Ho, 1995).
In ABA-deficient orange plants, an increase in stomatal opening on the fruit
caused higher fruit water loss than in wild-type oranges (Romero et al.,
2013). This also demonstrated the role of stomata in the water loss of fruits.
Therefore, fast handling after harvest is important in order to restrict water
loss and quality loss in fruits with higher stomatal densities.

6.4  Preharvest Conditions Leading to

Postharvest Problems via Stomata
There are several preharvest factors that can influence the postharvest per-
formance of horticultural crops (Sams, 1999; Lee and Kader, 2000; Herppich
et al., 2001; Torre and Fjeld, 2001; Tijskens et al., 2003; Rezaei Nejad and
van Meeteren, 2005; Hewett, 2006; In et al., 2007; Fonseca et al.,  2009;

Stom ata a n d P ostharvest P hy s i o l o g y

Moran  et  al., 2009; Islam et al., 2010; Fanourakis et al., 2011; Burchi and
Prisa, 2013; Tudela et al., 2013). During growth of the plants, the manage-
ment of environmental conditions such as relative humidity (RH), light, and
temperature can considerably influence postharvest performance of horti-
cultural products. Some of these conditions affect postharvest performance
via effects on the stomata (number, morphology, and closing ability).

6.4.1  Relative Humidity and Stomata Control

Commercial production of vegetables, cut flowers, and ornamental plants
under closed and semiclosed environments has tremendously increased
over the last few decades. In this kind of protected cultivation system,
often the RH inside the production facility is high because of energy sav-
ing (reduced ventilation and increased insulation) and transpiration by the
plants. RH is the ratio of the partial pressure of water vapor of the air to the
saturated water vapor pressure, expressed in percentages. Air saturates
when it holds water with its maximum capacity, which depends on tem-
perature. More moisture beyond this capacity would lead to condensation
of water vapor molecules to liquid water. By increasing the temperature,
the maximum water holding capacity of the air increases. Therefore, the
RH depends on the actual water vapor pressure of the air and air tempera-
ture. Vapor pressure deficit (VPD) is defined as the difference between
the saturation water vapor pressure and the actual water vapor pressure.
According to Fick’s first law, the diffusion rate of water vapor from a leaf
is linearly related to the difference in water vapor pressure inside and out-
side the leaf. Therefore, the humidity in plant environments is preferably
expressed as VPD.
Plants that have been produced under low-VPD conditions (very
humid) have often grown normally, but after harvest their stomata do not
function in a normal way anymore. Due to the prolonged exposure to low
VPD, a habituation process occurs in guard cells of the stomata, which
makes the stomata insensitive to stimuli that would normally provoke sto-
matal closure (stomatal malfunctioning) (Rezaei Nejad and van Meeteren,
2005; Aliniaeifard et al., 2014). As a result, stomata also stay open when
they encounter desiccation after harvest of the plants or plant products.
This disturbance in the normal functioning of the stomata due to previ-
ous exposure to low VPD therefore has horticultural consequences. For
example, as discussed before, growing rose plants at low-VPD conditions
often results in a decrease in vase life after harvest of the plants (Torre
et al., 2001). Low-VPD-grown plants often have a higher rate of water loss
than the moderate-VPD-grown plants; this is also the case during desic-
cation or hampered water uptake (postharvest stage) or when they are


exposed to high VPDs (Rezaei Nejad and van Meeteren, 2005; Rezaei
Nejad et al., 2006; Aliniaeifard and van Meeteren, 2013; Fanourakis et al.,
2013a; Aliniaeifard, 2014; Aliniaeifard et al., 2014). Such a transfer from
low to high VPD is common after vegetative propagation by leafy cuttings,
after in vitro propagation, and at the end of the cultivation period of orna-
mentals when plants or cut flowers are transferred to domestic conditions.
Although plants can be cultured in vitro on a large scale under low-VPD
conditions, in vitro–produced plants are usually susceptible to wilting
upon transfer to normal atmospheric VPDs (Brainerd and Fuchigami,
1982; Ghashghaie et al., 1992; Santamaria et al., 1993; Aguilar et al., 2000;
Hronková et al., 2003; Hazarika, 2006; Aracama et al., 2008; Khan et al.,
2010). This is also because of malfunctioning of the stomata in response to
a wide range of closing stimuli, such as darkness, abscisic acid (ABA), and
elevated calcium (Ca 2+) levels (Brainerd and Fuchigami, 1982; Ziv et al.,
1987; Santamaria et al., 1993). It has been shown that higher rates of water
loss after desiccation (postharvest phase) in the plants grown at low-VPD
conditions are mainly caused by the stomata, compared to the role of the
cuticle (Ziv et al., 1987; Santamaria and Kerstiens, 1994; Aliniaeifard and
van Meeteren, 2013; Fanourakis et al., 2013a).
As discussed in specific sections before, cultivation at low VPD not
only influences the vase life and visual appearance of cut flowers, but also
can influence the postharvest life and nutritional quality of leafy vegetables
via the stomata. Leaves of plants that lose their moisture more easily are
more vulnerable to losing vitamin than plants that are resistant to wilting
(Ezell and Wilcox, 1959; Lee and Kader, 2000).  How Does Preharvest Low VPD Affect

Postharvest Stomata Control?
Stomata react very fast to changes in the environmental conditions through
internal signals (Martin and Meidner, 1971; Wigger et al., 2002; Tallman,
2004; Neill et al., 2008; Kim et al., 2010; Hao et al., 2011; Hossain et al.,
2011). In natural conditions, plants continuously encounter changes in the
environment. Therefore, in order to survive, they should have the ability
to react fast to the changing environment. ABA is the main phytohormone
that acts as an internal signal in response to changes in environmental
conditions and triggers changes in various plant physiological and develop-
mental processes, resulting in adaptation to the stress conditions, including
stomatal closure (Aguilar et al., 2000; Chen et al., 2010; Lee and Luan, 2012).
In general, the ABA level is regulated by a balance between its biosynthesis
and its catabolism (Nambara and Marion-Poll, 2005; Lee and Luan, 2012).
The first step in the specific ABA production pathway is the synthesis of
violaxanthin through zeaxanthine epoxidase. Neoxanthin synthase and an
isomerase may be required for formation of cis-isomers of violaxanthin

Stom ata a n d P ostharvest P hy s i o l o g y

and neoxanthin. 9-cis-Epoxycarotenoid dioxygenases (NCEDs) cleave the

cis-xanthophylls to yield xanthoxin. It has been shown that NCED is the
rate-limiting factor in the ABA biosynthetic pathway (Qin and Zeevaart,
1999; Iuchi et al., 2001; Tan et al., 2003). Through a short-chain alcohol
dehydrogenase (ABA2), xanthoxin is then converted to abscisic aldehyde.
Finally, abscisic aldehyde oxidase (A AO3) catalyzes the last step of ABA
biosynthesis by oxidation of abscisic aldehyde into ABA (Nambara and
Marion-Poll, 2005).
The place in the plant of ABA biosynthesis and the place of its action
are still under debate (Christmann et al., 2005; Davies et al., 2005; Endo
et al., 2008; Jiang and Hartung, 2008; Melhorn et al., 2008). Following
drought stress, root-produced ABA acts as a signal between roots and
shoots (Holbrook et al., 2002; Wilkinson and Davies, 2002; Davies et al.,
2005; Jiang and Hartung, 2008). However, it has also been indicated that
synthesis of ABA in the shoot is a response to a long-distance hydraulic sig-
nal in xylem vessels due to low water potential in the soil (Holbrook et al.,
2002; Christmann et al., 2005, 2007, 2013). Nonetheless, it has been shown
that NCED3 is mainly expressed and localized in the vascular parenchyma
of leaves (Cheng et al., 2002; Endo et al., 2008). The expression of AAO3 in
the guard cells has also been reported (Koiwai et al., 2004; Nambara and
Marion-Poll, 2005; Melhorn et al., 2008). Moreover, the transient expres-
sion of NCED3 or AAO3 in guard cells promotes stomatal closure (Melhorn
et al., 2008). Bauer et al. (2012) showed that guard cells possess the entire
ABA biosynthetic pathway. Guard cells are able to autonomously synthesize
ABA, and there is a positive feedback loop for ABA production when they
have been exposed to high VPDs around the leaves (Bauer et al., 2012). It
has been suggested that there is no need for root–shoot transport of ABA
(Osakabe et al., 2013). Moreover, increasing VPD around the leaves of well-
watered plants resulted in a higher ABA level in the leaf (Rezaei Nejad and
van Meeteren, 2007, 2008).
ABA inactivates mainly through oxidation or conjugation processes.
Hydroxylation of ABA is the main process for ABA inactivation (Nambara
and Marion-Poll, 2005). Oxidation of ABA is catalyzed by 8’-hydroxylases
to form 8’-hydroxy ABA. In the next step, 8-hydroxy ABA spontaneously
isomerizes to phaseic acid (PA) and is further reduced to dihydrophaseic
acid (DPA) through an unknown reductase (Krochko et al., 1998; Cutler and
Krochko, 1999). Similar to PA, neophaseicacid (neoPA) can be formed from
hydroxy ABA through isomerization (Zhou et al., 2004). ABA 8-hydroxy-
lases are members of a CYP707A subfamily of cytochrome P450 monooxy-
genases (Kushiro et al., 2004; Saito et al., 2004). It has been reported that
exposure of plants to low VPD induces catabolism of ABA via CYP707As
genes (Okamoto et al., 2009).
Apart from the oxidative catabolic pathways, ABA can be inacti-
vated via conjugation with glucose to form its glucose ester (ABA-GE)


(Xu et al., 2002; Priest et al., 2006). ABA-GE is readily reversible but
not easily permeable to biomembranes and may function as a realizable
and transportable form of ABA (Dietz et al., 2000; Sauter et al., 2002).
When ABA is needed, ABA-GE is hydroxylated through β-glucosidase
to increase the active form of ABA (Lee et al., 2006). The activity of
β-glucosidase was decreased in rose plants that had been grown in low
VPD compared with its activity in moderate-VPD-grown plants. This
resulted in a higher ratio of ABA-GE to ABA (Arve et al., 2012). However,
it is still not clear whether the lower ABA level after long-term exposure
to low VPD is due to lower production or the higher catabolism of ABA.
It has been shown that exposure to different VPDs influences ABA in
the leaves of plants. In many plant species, such as spinach (Spinacia olera-
cea) (Zeevaart, 1974), rose (Arve et al., 2012; Giday et al., 2013a), Arabidopsis
(Okamoto et al., 2009), and spiderwort (Tradescantia virginiana) (Rezaei
Nejad and van Meeteren, 2007, 2008), the foliar [ABA] decreases as a result
of exposure to low-VPD conditions. In Tradescantia, 1 day after transfer-
ring moderate-VPD-grown plants to a low-VPD condition, the foliar [ABA]
decreased to the ABA level found in plants grown at low VPD. Reciprocal
transfer of moderate-VPD-grown plants back from low to moderate VPD
increased the foliar [ABA] again to its original level in moderate-VPD-
grown plants (Rezaei Nejad and van Meeteren, 2008). In rose plants, con-
trary to moderate-VPD-grown plants, exposure of low-VPD-grown plants
to darkness did not result in elevation of the foliar [ABA] (Arve et al., 2012;
Giday et al., 2013a). In Arabidopsis, the [ABA] decreased sharply even 1 h
after exposure to a low-VPD condition (Okamoto et al., 2009). Moreover,
in vitro–propagated plants that were produced under low-VPD conditions
were deficient in ABA (Hronková et al., 2003). Therefore, in the absence of
stresses, the foliar [ABA] depends on the VPD as well; decreasing the VPD
will result in a decrease in [ABA].
A foliar threshold level for ABA has been suggested in order to keep
the stomata responsive to closing stimuli. Since the foliar ABA level in mod-
erate-VPD-exposed plants is always higher than this threshold, their sto-
mata are always responsive. In different rose cultivars and also Arabidopsis
accessions, higher ABA levels than the threshold were still present after
low-VPD exposure; these genotypes maintained responsiveness to closing
stimuli in the postharvest phase (Giday et al., 2013a; Aliniaeifard and van
Meeteren, 2014). In genotypes that showed malfunctioning of stomata after
a 4-day exposure to low VPD, a daily spray of ABA during the 4-day expo-
sure to low VPD resulted in maintaining the stomatal closing response to
closing stimuli in Arabidopsis and bean (Aliniaeifard and van Meeteren,
2013, 2014; Aliniaeifard, 2014; Aliniaeifard et al., 2014). This confirmed that
a long-term low ABA level results in loss of stomatal closing ability after-
wards. Changes in the signaling pathway inside the guard cells of stomata
due to long-term exposure to low-VPD conditions are reviewed in detail by

Stom ata a n d P ostharvest P hy s i o l o g y

Aliniaeifard and van Meeteren (2013). They proposed that alterations in

signaling pathways due to a long-term low transpiration rate under long-
term exposure to environmental conditions, especially low VPD, lead to
depletion of ABA, Ca 2+, and H 2O2 in the guard cells, as well as depletion of
extracellular ABA and Ca 2+. This will be accompanied by a low sensitivity
for ABA, due to a strong negative regulation of the ABA signaling path-
way by the type 2C protein phosphatases ABI1 and ABI2 and a low positive
regulation of the ABA signaling pathway by OST1. These effects will be
strengthened by a low sensitivity of the anion channel SLAC1 for Ca 2+. This
coincidence in changes of Ca 2+, ABA receptors, and positive and negative
regulators of ABA signaling is proposed as an explanation for persistency
of the stomatal malfunctioning induced by long-term exposure to low VPD.  Induction of Stomata Morphological

Changes by Low VPD
Besides stomatal closing ability, stomatal size and density can also be
influenced by VPD (Fordham et al., 2001; Tricker et al., 2012). Rose,
Tradescantia, and bean plants that developed their leaves in low-VPD
conditions are characterized by large stomata and a wide aperture area
(Figure  6.3) (Torre et al., 2003; Rezaei Nejad and van Meeteren, 2005;
Fanourakis et al., 2011, 2013a; Aliniaeifard et al., 2014). A question arises:
Are stomatal morphological alterations involved in the decreased ­stomatal
closing ­
ability of low-VPD-grown plants after harvest? A connection
between stomatal function and structural features for various species
has previously been suggested (Franks and Farquhar, 2007; Doheny-
Adams et al., 2012; Drake et al., 2013; Giday et al., 2013b). Because of the
higher ratio of a guard cell’s membrane surface to its volume, species with
smaller stomata may respond faster than species with larger stomata. On


100 μm 100 μm

Figure 6.3  Stomatal size and density in low (L)- and moderate (M)-VPD-
grown Vicia faba plants.


the other hand, smaller stomata are usually associated with higher stoma-
tal density per leaf area (Hetherington and Woodward, 2003). To optimize
the trade-off between carbon gain and transpirational water loss, these
characteristics (smaller stomata and higher stomatal density) allow the
leaf to attain high stomatal conductance under favorable conditions, and
to promptly reduce stomatal conductance when conditions are unfavor-
able, which help the plant cope with stress conditions (Xu and Zhou, 2008;
Doheny-Adams et al., 2012). Using five closely related species of the genus
Banksia, it has been demonstrated that the rate of stomatal response was
negatively correlated with stomatal size (Drake et al., 2013). It has been
indicated that smaller stomata require less leaf drying to close and that
plants’ stomatal size underlays much of the variation in the regulation of
transpiration upon desiccation (postharvest phase) after growth of the
plants in low and moderate VPDs (Franks and Farquhar, 2007; Doheny-
Adams et al., 2012; Drake et al., 2013; Giday et al., 2013b).
In cut rose flowers, although changes in stomatal length had no
influence on stomatal functionality, anatomical features per se represent a
significant and direct contribution to the increased water loss in the post-
harvest phase of the roses (Fanourakis et al., 2013a). Higher stomatal den-
sities enlarge the stomatal pore area per leaf area, leading to an increase
in the transpiration rate in low-VPD-grown plants. However, in faba bean,
stomatal density decreased as a result of growing them at low VPD. On
the other hand, in faba bean, stomata of low-VPD-grown plants were
larger for all guard cell dimensions than those of moderate-VPD-grown
plants (Aliniaeifard et al., 2014). Therefore, high stomatal conductance
in low-VPD-grown plants can be attributed to their larger stomata with
wider pore area in comparison with the stomata in m ­ oderate-VPD-grown
plants. In the previous studies, investigating the impact of stomatal mor-
phological traits on stomatal functionality, the stomatal responsiveness
of different species was investigated or plants from one or several geno-
types were exposed to contrasting environments, which usually induced
changes in both stomatal morphology and responsiveness (Hetherington
and Woodward, 2003; Torre et al., 2003; Franks and Farquhar, 2007; Drake
et al., 2013; Fanourakis et al., 2013a; Giday et al., 2013b). Analyzing the
importance of the stomatal morphological traits on the stomatal function-
ality, not only in moderate and low-VPD-grown plants, but also in plants
that had developed their leaves in moderate VPD and thereafter trans-
ferred for 1–4 days to low VPD, confirmed that stomata ­morphological
traits are not always determinant of stomatal functionality: the morphol-
ogy of stomata in plants that had developed their leaves at moderate VPD
and were then transferred for 4 days to low VPD was more similar to that
of ­moderate-VPD-grown plants, while their response to ABA and des-
iccation was similar to the stomatal response of low-VPD-grown plants
(Aliniaeifard et al., 2014).

Stom ata a n d P ostharvest P hy s i o l o g y

It has been suggested that changes in the signaling related to ABA

are the determining factor in the decreased closing ability of stomata and,
as a result, decreased vase or shelf life of horticultural crops after long-term
exposure to low VPD (Aliniaeifard and van Meeteren, 2013; Aliniaeifard,
2014; Aliniaeifard et al., 2014). It can be concluded that production of flow-
ers and vegetables in low VPD renders the stomata incapable of a suitable
response to closing stimuli afterwards. In the flowers, their rate of water
uptake becomes lower than their transpiration rate, resulting in a shorter
vase life due to a negative water balance, and leafy vegetables have a
reduced capacity to keep water after harvest, resulting in a loss of textural
and nutritional quality.

6.4.2 Temperature
As discussed in Section 6.2.1, stomata are involved in controlling plant
t emperature through the transpiration rate. Therefore, it could be
expected that stomata will open more with increasing temperatures to
prevent overheating of the plant. However, the response of stomata to tem-
perature is not constant for different temperatures, and usually it is species
dependent. There are contradictory reports regarding stomatal response
to different temperatures. Both stomatal closure and stomatal opening
have been reported as a result of an increase in temperature (Schulze
et al., 1973; Wilkinson et al., 2001). Likely this is because stomata opening
responds to many factors; among these factors are the CO2 concentration
in the stomatal cavity and the water potential of the leaves. These param-
eters are respectively affected by the rate of photosynthesis and by water
uptake and water loss. These processes are also largely influenced by
temperature. Schulze et al. (1973) showed for five species (Zygophyllum
dumosum, Artemisia herba-alba, Hamada scoparia, Reaumurian egevensis,
Prunus armeniaca) that stomata opened more in response to a gradual
increase in temperature (from 31.6°C to 39.8°C) at low water stress. This
response was independent of VPD. At high plant water stress, however,
the stomatal response was reversed; that is, the stomata closed when the
temperature was gradually increased. In leaf disks of legumes floating on
water, stomata opened in darkness by increasing the temperature from
20°C to 45°C (Feller, 2006; Reynolds-Henne et al., 2010), even when ABA
was applied. By using epidermal strips floating on water, the direct effect
of temperature on guard cells was studied, excluding the effects of pho-
tosynthesis and water loss (Rogers et al., 1979). They showed that stoma-
tal aperture increased by an increase of temperature from 10°C to 40°C,
while at 45°C the aperture was less than those obtained at 35°C and 40°C.
In the experiments mentioned, there was a fast recovery of stomata to its
original apertures when temperature was decreased again. This indicates


that large aftereffects of cultivation temperature on postharvest behavior

are not expected.
There are some reports that stomatal density decreases with increas-
ing temperature (Beerling and Chaloner, 1993). Stomatal density of
Dodonaea, produced as cut foliage branches, showed a strong correlation
with seasonal average monthly day temperature. The cut branches showed
also strong seasonal variations in longevity, wilting after 1 week in winter,
while displaying a vase life of 3 weeks in summer. On day 1 of vase life, the
branch water status was positively correlated with vessel, stomata, and tri-
chome densities, and negatively correlated with vessel member length and
leaf thickness. However, on day 16, the branch water status was negatively
correlated only with vessel member length and diameter, implying a differ-
ent relative importance of anatomical parameters for surviving water stress
in the vase. The limited part that stomata number plays in the vase life of
cut flowers was discussed in Section 6.2.
Mostly, horticultural products are cooled down after harvest. Rapid
stomatal closure can be induced by exposure to low temperatures. However,
there is variation in the stomatal closure response between species after
exposure to low temperatures. For example, in response to cold (7°C), sto-
mata of Commelina communis are closed within 1 hour of transferring the
plants from 27°C to 7°C, whereas the stomata of tobacco (Nicotiana rustica)
are not capable of closure, which leads to uncontrolled water loss and wilt-
ing of the leaves after 5 hours of the transfer to low temperature (Wilkinson
et al., 2001). The rapid low-temperature-induced stomatal closure is likely
to be the result of a low-temperature increase in guard cell plasma mem-
brane calcium channel activity. Chilling-sensitive plants, such as tobacco,
do not show this increase in calcium uptake at low temperature (Wilkinson
et al., 2001). It is also possible that in chilling-sensitive produce, the mem-
branes of epidermal cells lose their semipermeability characteristics, and
thus their turgor, before guard cells lose them. In that case, the stomata
will also open. It can be concluded that the effect of temperature during
the postharvest phase on the function of stomata, as well as on keeping the
quality of plant produce, is highly species dependent. Both low and high
temperatures can influence the proper function of stomata and, as a result,
influence the quality of the products. The effect of preharvest temperature
on the role of stomata in keeping quality is very limited.

6.4.3 Light
Greenhouse crop production frequently makes use of supplementary light-
ing to improve plant productivity in periods of the year when natural irradi-
ance is low; on some occasions, continuous lighting is used (Mortensen
and Fjeld, 1998; Dodd et al., 2005; Pettersen et al., 2006; Velez-Ramirez

Stom ata a n d P ostharvest P hy s i o l o g y

et al., 2011). For example, in pot roses, extending the lighting period from
18 to 24 h per day shortened the time to flowering by 12% and the num-
ber of flowers increased by 34% (Pettersen et al., 2007). Although grow-
ing plants under continuous light has several advantages (e.g., reduction
of powdery mildew), it adversely influences the postharvest life of the cut
flowers (Mortensen and Fjeld, 1998; Mortensen and Gislerød, 1999, 2011)
due to poor stomatal closure under conditions of decreasing leaf water
potential and turgor (Slootweg and van Meeteren, 1991; Arve et al., 2012).
The vase life of the roses decreased by using supplemental lighting. The
vase life depends on the duration of exposure to light. The longer the light-
ing period during growth, the shorter the vase life afterwards (Mortensen
and Fjeld, 1998; Pettersen et al., 2006; Mortensen and Gislerød, 1999, 2011;
Arve et al., 2012). Extending the lighting period from 16 to 20 h decreased
the postharvest life in lemon balm, while it is not affected in basil; however,
24 h exposure to light resulted in decreased postharvest life in both herbs
(Islam et al., 2010). It has been shown that by decreasing the RH in the
growing units when plants are grown under continuous light, it is possible
to reduce the disability of stomata to close and, as a result, increase the vase
life of cut flowers (Mortensen and Fjeld, 1998; Mortensen and Gislerød,
1999, 2011; Pettersen et al., 2006; Arve et al., 2012). However, even in mod-
erate RH conditions, exposure to continuous light results in decreased vase
life of cut roses (Mortensen and Fjeld, 1998).
Exposure to light not only influences the sugar content through pho-
tosynthesis, but also can induce sugar accumulation because of osmotic
adjustment in the cells. For example, higher sugar levels have been found
in the rinds of mandarin fruits positioned outside of the tree canopy fac-
ing toward sunlight. Sugar accumulated (osmotic adjustment) in the rinds
of sunlight-exposed fruits because of dehydration of the rind’s cells due
to high transpiration via stomata of sun-exposed fruit surfaces (Yakushiji
et  al., 1998; Magwaza et al., 2013). Accumulation of sugar and other bio-
chemicals in the rinds of sun-exposed fruit during the growing season
improves their postharvest performance and decreases susceptibility to
rind breakdown (Magwaza et al., 2013).
The amount of healthy compounds of a plant can be influenced by
the lighting period and light intensity. Although light has no direct role
in the production of ascorbic acid (vitamin C), the duration and intensity
of light can influence the production of ascorbic acid in the leaf (Lee and
Kader, 2000). In general, the amount of ascorbic acid increases by light
(intensity and duration) and decreases in the dark period (Lee and Kader,
2000). On the other hand, plants that have been produced in a prolonged
lighting period could lose their vitamin C more easily after harvest because
of higher water loss via stomata after harvest; therefore, there should be
a balance for the lighting period for producing vitamin C before harvest
and preventing loss of vitamin C after harvest of leafy vegetables. However,


research in which the effect of light during cultivation on vitamin C content

during the pre- and postharvest phases is lacking. Also, the amount of other
secondary metabolites can be affected by light. For example, linalyl acetate
is one of the compounds in the essential oil of oregano plants with anti-
inflammatory activity (Peana et al., 2002). It has been found that growing
oregano plants in full light resulted in a decreased amount of linalyl acetate
due to more open stomata and higher transpiration rate. On the other hand,
growing plants in pots under 50% shade resulted in a higher percentage of
linalyl acetate in the essential oil (Tibaldi et al., 2011).

6.5  Stomata and Tolerance to Postharvest

Diseases and Physiological Disorders
Postharvest diseases reduce the quantity and quality of horticultural prod-
ucts. Although some diseases may not lower produce saleability, they still
reduce product value. Losses in value of the product through postharvest
diseases may occur at any time from harvest to consumption (Coates and
Johnson, 1997). The entrance of pathogens to plant tissues is the first and
critical stage during the infection process. Some pathogenic and sapro-
phytic microorganisms are able to survive and reduplicate on the leaf sur-
face without infection (Melotto et al., 2008). Although many of pathogenic
fungi can penetrate the cuticle and epidermis by different means (Mendgen
et al., 1996), some fungal diseases and many pathogenic bacteria are unable
to directly penetrate the host cuticle. The cuticle plays a key role in the
survival of plants against biotic and abiotic stresses. The primary physi-
ological role of the plant cuticle is to seal the tissue against a relatively dry
atmosphere, preventing desiccation by minimizing water loss (Mintz-Oron
et al., 2008; Riederer and Muller, 2008); the position of stomata in the inter-
face between the surrounding environment and leaf internal tissues adds
another role for this microscopic pore. It provides a route for endophytic
colonization of pathogens in the plant tissues; therefore, plants also have
established a mechanism to regulate stomatal aperture not only in response
to a range of environmental factors, but also in response to pathogens
(Melotto et al., 2008; Gudesblat et al., 2009). Both stomatal density and
size are important factors determining the infection of products by patho-
gens. Asiatic citrus canker caused by Xanthomonas citri subsp. citri seri-
ously affects the citrus industry in Asia. It causes formation of pustule-like
necrotic lesions on the fruit, and it highly decreases fruit quality. Stomatal
characteristics such as stomatal density, size, and closing response will
influence the magnitude of infection of fruit internal tissues. Stomatal char-
acteristics can be considered the main factor influencing the resistance
magnitude of citrus plants to the bacteria (Wang et al., 2011a). Therefore,

Stom ata a n d P ostharvest P hy s i o l o g y

the presence of more stomata and a larger stomatal aperture area in the fruit
surface could increase the susceptibility to bacteria. Wang et al. (2011a)
found that under the same experimental conditions, ‘Meiwa’ kumquat has a
lower stomatal density and smaller stomatal openings than ‘Newhall’ navel
orange. These stomatal characteristics of ‘Meiwa’ make this cultivar better
resistant to canker than ‘Newhall’. ‘Pinalate’ (Citrus sinensis L. Osbeck) is a
yellow ABA-deficient mutant derived from the orange ‘Navelate’; therefore,
its stomata will close less. ‘Pinalate’ has a higher transpiration rate, and due
to the impaired stomatal functioning, it is more susceptible to pathogenic
infections (Alférez et al., 2005).
Since stomata are the entrance site of many bacterial and fungal
diseases into leaves or fruits of horticultural products, the immunity of a
product to pathogen virulence in many cases depends on the function of
the stomata (Zeng et al., 2010). Many postharvest treatments are efficient
when they restrict the entrance of pathogens into the products. For exam-
ple, heat treatments such as hot water dips, vapor heat, short hot water
rinse, and brushing, causing melting of the wax layer in the surface of the
fruits, and therefore covering of the stomata, provide a mechanical barrier
for the pathogens, restricting the invasion of pathogens into the internal
tissues (Porat et al., 2000; Schirra et al., 2000; Waller et al., 2001; Mulas
and Schirra, 2007). On some occasions, exogenous wax is applied in order
to extend the quality of the fruits. The amount of wax applied on the fruit is
important to prevent pathogen infection. Low-coating loads result in some
uncoated surface areas with bare stomata, while sufficient wax loads cover
all the stomata and restrict pathogen infections (Njombolwana et al., 2013).
Salicylic acid is one of the plant hormones known to be involved in sensing
the pathogenic invasion. Salicylic acid–induced stomatal closure is one of
the cellular pathways that has been suggested after infection by pathogens
(Lu and Chen, 2005; Melotto et al., 2006; Khokon et al., 2011). It has been
shown that inducing stomatal closure by salicylic acid improves the plant
capacity to cope with diseases. For example, in lily, salicylic acid treatments
lead to induction of stomatal closure, causing more tolerance of lily plants to
Botrytis elliptica (Lu and Chen, 2005).
In some products, monitoring stomatal conductance can be used
as a nondestructive index for storability and quality status. Controlling
water loss from the leaves (especially from the stomata) can be consid-
ered an important way to minimize quality problems and extend the
postharvest life in crops such as rose cut flowers and kalanchoe cuttings
(Bredmose and Nielsen, 2009; Fanourakis et al., 2013a, 2013b). Stomatal
density in tomato fruits is very low, whereas the number of stomata in
cucumber is relatively high (Table 6.2). Higher stomatal densities posi-
tively correlate with higher water loss from the fruits. Measuring the
amount of water loss immediately after harvest showed 20 times higher


water loss for cucumber fruits than tomato fruits (Adams and Ho, 1995).
Therefore, fast handling after harvest is important in order to restrict
water loss and quality loss in crops with higher stomatal densities. High
humidity during growth decreases movement of calcium to the leaves in
cucumber and tomato plants. However, due to the higher stomatal den-
sity in the fruits of cucumber than in tomato fruits, a higher level of cal-
cium accumulates in the cucumber fruits (Adams and Ho, 1995). This
confirmed that transpiration by fruits plays a pivotal role in determining
calcium levels in the fruits and susceptibility to physiological disorders
related to calcium deficiency.

6.6  Postharvest Treatments and Stomata

In many horticultural products, the stomata are functional after harvest,
and their function is important for postharvest quality (Johnson and Brun,
1966; Moreshet and Green, 1980; Blanke and Lenz, 1989; Burg, 2004).
Storage of fresh-cut Romaine lettuce in light conditions increases the num-
ber of open stomata and results in increased water loss and increased occur-
rences of leaf browning (Martínez-Sánchez et al., 2011). Stomatal opening
in Chinese kale positively correlates with loss of fresh weight. However,
vitamin C content in light-stored Chinese kale leaves was higher than in
dark-stored leaves (Noichinda et al., 2007).
Postharvest potassium silicate application covers fruit stomata with a
silicate layer, which positively delays fruit weight loss by keeping fruit mois-
ture (Tesfay et al., 2011). In some horticultural products, postharvest heat
treatments decrease water loss from the product instead of increasing it.
This effect of postharvest heat treatments is due to their effects on the melt-
ing of cuticular wax, which causes coverage of stomata and microcracks;
therefore, heat treatments are able to reduce water loss by decreasing tran-
spiration (Schirra et al., 2000). For example, treatment of Valencia oranges
by 45°C water for 42 min keeps the firmness of the fruit and considerably
decreases water loss. But immersion of fruits in 53°C water for 12 minutes
increases the water loss from the fruits (Williams et al., 1994; Tang et al.,
2007). However, by shortening the time of hot water immersion, it would
be possible to use hotter water to control water loss. For example, treat-
ing orange fruits at 56°C for 20 s also melts epicuticular waxes, leading to
covered and sealed stomata and cracks on the fruit without occurrences of
other related problems, such as fruit weight loss, surface damage, or inter-
nal quality loss (Porat et al., 2000).
Postharvest pathogens are one of the items considered for loss of
quality during storage of the products. Since the main ports for entrance
of pathogenic disease are stomata, covering of stomata by wax has the
potential to reduce the incidence of postharvest diseases by providing a

Stom ata a n d P ostharvest P hy s i o l o g y

physical barrier against pathogens (Schirra et al., 2000; Li et al., 2013).

Blocking stomata and other cracks by treating orange fruits at 56°C for
20 s, also restricts occurrences of Penicillium digitatum (Porat et al., 2000).
Although ethylene can induce some quality problems in the postharvest
stage (hastening ripening or senescence), treatment with ethylene forms
new waxes on the stored fruits that cover stomata, cracks, or areas lacking
wax and provides mechanical barriers to P. digitatum penetration (Cajuste
et al., 2010). As another role in postharvest treatments, the presence of
stomata facilitates the absorption of 1-methylcyclopropene (1-MCP) into
product tissues (Dong et al., 2013), a gas often used to inhibit senescence
as it blocks ethylene receptors.
Storage of products in low-pressure-type storage induces stomatal
malfunctioning in response to darkness (Laurin et al., 2006). The problem
with stomatal malfunctioning worsens when the product is stored at low
pressure for a long time period. Under such conditions, it has been reported
that the marketable period shortens and the weight loss of cucumber
increases by 5% (Laurin et al., 2006). Similarly, in grape berries, 20% of the
stomata stay fully or partially open at darkness (Blanke and Leyhe, 1988).
Kirk et al. (1986) reported that long-term storage of fresh hibiscus cuttings
in low-pressure storage induces stomatal malfunctioning in response to
several closing stimuli, such as high carbon dioxide and low RH.
The stomatal closing effect of ABA has been used to control water loss in
order to extend the postharvest life of cut flowers and potted plants (Cornish
et al., 1985; Pompodakis et al., 2004; Shimizu-Yumoto and Ichimura, 2009;
Reid and Jiang, 2012). Since ABA is an expensive compound, some stud-
ies have tried to find a way to increase the endogenous ABA level with less
expensive methods. In some studies, NaCl or osmotic stress was applied to
the plants to close the stomata in order to extend their postharvest life (Reid
and Jiang, 2012; Giday et al., 2014). S-Abscisic acid (S-ABA), the biologically
active form of ABA produced through microbial fermentation, is commer-
cially used for extending postharvest life of products. However, in some spe-
cies, some damage to the flower and shoot has been reported by using this
compound (Runkle et al., 2007; Waterland et al., 2010). It seems that using a
high concentration of S-ABA is problematic, while using low concentrations
can extend postharvest life without negative effects (Reid and Jiang, 2012).
However, senescence can be enhanced by application of ABA (Halevy et al.,
1974; Ferrante et al., 2004; Hunter et al., 2004).

6.7  Ethylene, Stomata, and Senescence

To integrate all the plant responses to the environment, phytohormones play
an important role, as well as the interplay between them. The role of ABA
as the main phytohormone reducing transpiration by provoking stomatal


closure is mentioned above. There are contradictory reports regarding sto-

matal responses to ethylene. Both induction of stomatal closure (Vitagliano
and Hoad, 1978; Pallas and Kays, 1982; Taylor and Gunderson, 1986, 1988;
Tissera and Ayres, 1986; Desikan et al., 2005, 2006) and promotion of sto-
matal opening (Frommhold, 1982; Merritt et al., 2001; She and Song, 2012;
Song et al., 2012) have been reported for ethylene. However, in some stud-
ies, no effect of ethylene on stomatal opening and closure has been reported
(Pallaghy and Raschke, 1972; Bradford, 1983; Madhavan et al., 1983). The
interplay between ethylene and other phytohormones is likely to be determi-
nant in the regulation of stomatal movements by ethylene (Pallas and Kays,
1982; Merritt et al., 2001; Tanaka et al., 2005, 2006; Acharya and Assmann,
2009; Chen et al., 2013). It has been reported that induction of stomatal open-
ing or inhibition of stomatal closure by auxins and cytokinins is through their
promotional effect on ethylene production (Merritt et al., 2001; Tanaka et al.,
2006). It seems that stomata close in response to ethylene in the absence of
ABA and open in the presence of ABA (Desikan et al., 2006; Tanaka et al.,
2006). It has been shown that ethylene or ACC (the precursor of ethylene)
can prevent ABA-induced stomatal closure (Tanaka et al., 2005). Application
of ethylene increases stomatal aperture of wild-type Arabidopsis exposed to
drought conditions (Tanaka et al., 2005). It is likely that ethylene acts on the
late steps of the ABA-induced stomatal closure (Tanaka et al., 2005). The
interaction between AtERF7 (which is a member of the ethylene-responsive
binding factor) and PKS3 (a Ser/Thr protein kinase that interacts with ABI2
and, to some extent, ABI1) (Guo et al., 2002) decreases the guard cells’ sen-
sitivity to ABA and increases water loss through transpiration (Song et al.,
2005). For induction of stomatal closure by ethylene, it seems that ethyl-
ene close the stomata through an ABA-independent pathway. The ethylene
receptor ETR1 mediates H 2O2 signaling in guard cells and may be a sensor
for H 2O2 perception in guard cells (Desikan et al., 2005). So ETR1 func-
tions as the site of ethylene and H 2O2 cross talk, leading to stomatal closure
(Desikan et al., 2005).
Usually after harvesting, horticultural products are stored in dark-
ness and face water stress and high CO2 concentrations during storage. It
has been shown that water stress and high CO2 concentrations can induce
ethylene production. Stomatal responses are sensitive to darkness, water
stress, and CO2. As discussed before, it seems that the effect of ethylene
on stomatal closure highly depends on other factors, such as the presence
of other stresses and its interaction with other phytohormones. In order to
keep the quality of a product during storage, it is important to know the spe-
cies-specific stomatal responses of the product to CO2 and ethylene. Usually
the stomata of harvested leaves close instantaneously or within 10–35 min
because of harvesting procedures, and this considerably increases their
internal ethylene concentrations (Burg, 2004). On the other hand, CO2 can
inhibit the enhancing effect of ethylene on senescence (Aharoni et al., 1979;

Stom ata a n d P ostharvest P hy s i o l o g y

Sisler and Wood, 1988). The dual effect of high CO2 during storage is pre-
sented in Figure 6.4. At atmospheric pressure, closure of stomata occurs
during storage and transport in darkness, but using low-pressure (LP)
storage encourages stomatal opening even in darkness (Kirk and Skytt
Anderson, 1985; Kirk et al., 1986; Veierskov and Kirk, 1986). This low sto-
matal resistance at LP results in improved gas diffusion (such as ethylene)
and considerably improves intercellular ventilation. In Valencia orange with
open stomata, the transpirational resistance is 13.5 scm−1 (Moreshet and
Green, 1980). In the fruits, the stomatal resistance to ethylene transport is
23 scm−1. Due to stomatal closure during harvest, the resistance to ethyl-
ene transport increases to 6886 scm−1 in the orange fruits (Ben Yehoshua
et al., 1979). Moreover, fruit’s cuticular resistance to ethylene transport is
approximately 10-fold higher than its diffusive resistance to ethylene of
the stomata. Under the situation of closed stomata, the internal ethylene
concentration largely increases. Therefore, in order to prevent detrimental
effects of ethylene, it is vital to decrease the internal ethylene concentration
by some means, such as vacuum extraction (Burg, 2004).
Detaching organs from the mother plants and placing them in dark-
ness can promote leaf senescence in many plant species. A relationship
between stomatal aperture and senescence has been suggested (Thimann

High CO2 Darkness Desiccation

Stomatal closure

High internal harvest
ethylene level

(quality loss)

Figure 6.4  Diagram describing the effect of stomatal closing stimuli

­prevalent after harvest on the product quality.


and Satler, 1979). Senescence can be accelerated in darkness and slowed

down in light. The preventive effect of light on senescence is partly due to
its effect on stomatal opening, as suggested by the finding that promotion of
stomatal opening in darkness (using cytokinins, fusicoccin, etc.) postpones
senescence (in a similar way as light) (Thimann and Satler, 1979; Thimann,
1985). Cytokinins induce stomatal opening and delay senescence (Gan and
Amasino, 1995; Wingler et al., 1998; Song et al., 2006; Tanaka et al., 2006).
On the other hand, it has been shown that senescence can be accelerated by
ethylene. As discussed before, there is a controversy in reports regarding the
role of ethylene on stomatal closure. The rate of senescence level correlates
positively with the ABA level. Since antagonistic effects of cytokinins on ABA
have been proved, this can explain the preventive effects of cytokinins on
senescence (Thimann, 1985; Pospíšilová et al., 2000; Wang et al., 2011b).
In the flowers (petals) also, ABA regulates the process of senescence.
The level of ABA increases as senescence of petals starts. Also, a higher
ABA level has been shown in short-lived rose genotypes than in long-lived
genotypes (Mayak et al., 1972; Swanson et al., 1975). Confirming the role
of ABA in senescence, it has been reported that exogenous application of
ABA accelerates senescence in carnations (Mayak and Dilley, 1976) and
roses (Mayak et al., 1972).
The effect of the environment on leaf senescence is local, and it dif-
fers from young leaves to old leaves (leaf age dependency) (Weaver and
Amasino, 2001). It has been shown that the problem of low-VPD-induced
stomatal malfunctioning increases by leaf age; after exposure to low VPD,
stomata of old leaves are less responsive to desiccation than the stomata
of younger leaves (Rezaei Nejad and van Meeteren, 2007). In Arabidopsis,
after desiccation the rate of water loss increases by the leaf age; the expres-
sions of Senescence-Associated Gene113 (SAG113) and AtNAP (gene encod-
ing for a NAC family transcription factor, AtNAP) are co-induced during
leaf senescence (Zhang and Gan, 2012; Zhang et al., 2012). SAG113, a gene
that encodes a protein phosphatase belonging to the PP2C family, functions
as a negative regulator of the ABA signaling pathway and prevents stomatal
closure in response to closing stimuli such as ABA and desiccation (Zhang
et al., 2012). The AtNAP transcription factor physically interacts with the
promoter region of SAG113 and promotes its expression at the transcrip-
tional level (Zhang and Gan, 2012) and keeps the stomata less functional to
closing stimuli.

6.8  Conclusions and Future Perspectives

Climatic conditions in both protected and open-field cultivation can influ-
ence postharvest performance of horticultural products. During posthar-
vest handling and storage, many parameters, such as species, variety, light,

Stom ata a n d P ostharvest P hy s i o l o g y

humidity, temperature, radiation exposure, microbial load, presence of

pests, packaging, and chemical treatments, considerably change physical
and biochemical properties and the rate of deterioration of horticultural
products. Sometimes, growing plants in certain conditions results in the
optimum growth of the plants, but after harvest, several problems for the
product occur that lead to decreased shelf or vase life and, in the worse situ-
ations, deterioration of horticultural products.
Stomata are the main ports connecting internal leaf space to the out-
side environment. They can play an important role in connecting prehar-
vest factors to postharvest quality changes; this possible linkage function
has hardly been investigated.

Aarts, J.F.T. 1957. Over de houdbaarheid van snijbloemen [On the keepabil-
ity of cut flowers]). Mediterrenean Landbouwhogeschool Wageningen 57,
Acharya, B.R., and Assmann, S.M. 2009. Hormone interactions in stomatal
function. Plant Molecular Biology 69, 451–462.
Adams, P., and Ho, L. 1995. Differential effects of salinity and humidity on
growth and Ca status of tomato and cucumber grown in hydroponic
culture. Acta Horticulturae 401, 357–364
Agüero, M.V., Ponce, A.G., Moreira, M.R., and Roura, S.I. 2011. Lettuce qual-
ity loss under conditions that favor the wilting phenomenon. Postharvest
Biology and Technology 59, 124–131.
Aguilar, M.L., Espadas, F.L., Coello, J., Maust, B.E., Trejo, C., Robert, M.L.,
and Santamaría, J.M. 2000. The role of abscisic acid in controlling leaf
water loss, survival and growth of micropropagated Tagetes erecta
plants when transferred directly to the field. Journal of Experimental
Botany 51, 1861–1866.
Aharoni, N., Anderson, J.D., and Lieberman, M. 1979. Production and action
of ethylene in senescing leaf discs effect of indoleacetic acid, kinetin,
silver ion, and carbon dioxide. Plant Physiology 64, 805–809.
Alférez, F., Sala, J.M., Sanchez-Ballesta, M.T., Mulas, M., Lafuente, M.T., and
Zacarias, L. 2005. A comparative study of the postharvest performance
of an ABA-deficient mutant of oranges. I. Physiological and quality
aspects. Postharvest Biology and Technology 37, 222–231.
Aliniaeifard, S. 2014. Signal transduction pathways in guard cells after
prolonged exposure to low vapour pressure deficits. PhD thesis,
Wageningen University.
Aliniaeifard, S., and van Meeteren, U. 2013. Can prolonged exposure to
low VPD disturb the ABA signaling in stomatal guard cells? Journal of
Experimental Botany 64, 3551–3566.


Aliniaeifard, S., and van Meeteren, U. 2014. Natural variation in stomatal

response to closing stimuli among Arabidopsis thaliana accessions after
exposure to low VPD as a tool to recognise the mechanism of disturbed
stomatal functioning. Journal of Experimental Botany, doi: 10.1093/jxb/
Aliniaeifard, S., Malcolm Matamoros, P., and van Meeteren, U. 2014. Stomatal
malfunctioning under low VPD conditions: Induced by morphological
and anatomical or by signaling alterations? Physiologia Plantarum 152,
Amjad, M., Akhtar, J., Anwar-ul-Haq, M., Yang, A., Akhtar, S.S., and Jacobsen,
S.E. 2014. Integrating role of ethylene and ABA in tomato plants adapta-
tion to salt stress. Scientia Horticulturae 172, 109–116.
Antoni, R., Rodriguez, L., Gonzalez-Guzman, M., Pizzio, G.A., and Rodriguez,
P.L. 2011. News on ABA transport, protein degradation, and ABFs/
WRKYs in ABA signaling. Current Opinion in Plant Biology 14, 547–553.
Aracama, C.V., Kane, M.E., Wilson, S.B., and Philman, N.L. 2008. Comparative
growth, morphology, and anatomy of easy- and difficult-to-acclimatize
sea oats (Uniola paniculata) genotypes during in vitro culture and ex
vitro acclimatization. Journal of the American Society for Horticultural
Science 133, 830–843.
Arve, L.E., Terfa, M.T., Gislerød, H.R., Olsen, J.E., and Torre, S. 2012. High
relative air humidity and continuous light reduce stomata functional-
ity by affecting the ABA regulation in rose leaves. Plant, Cell and
Environment 36, 382–392.
Atkins, C.A., Kuo, J., Pate, J.S., Flinn, A.M., and Steele, T.W. 1977.
Photosynthetic pod wall of pea (Pisum sativum L.) distribution of car-
bon dioxide-fixing enzymes in relation to pod structure. Plant Physiology
60, 779–786.
Azad, A.K., Sawa, Y., Ishikawa, T., and Shibata, H. 2007. Temperature-dependent
stomatal movement in tulip petals controls water transpiration during
flower opening and closing. Annals of Applied Biology 150, 81–87.
Bakker, J.C. 1991. Effects of humidity on stomatal density and its relation to
leaf conductance. Scientia Horticulturae 48, 205–212.
Barbieri, G., Vallone, S., Orsini, F., Paradiso, R., De Pascale, S., Negre-
Zakharov, F., and Maggio, A. 2012. Stomatal density and metabolic
determinants mediate salt stress adaptation and water use efficiency in
basil (Ocimum basilicum L.). Journal of Plant Physiology 169, 1737–1746.
Barth, M.M., Perry, A.K., Schmidt, S.J., and Klein, B.P. 1992. Misting affects
market quality and enzyme activity of broccoli during retail storage.
Journal of Food Science 57, 954–957.
Bauer, H., Ache, P., Lautner, S., Fromm, J., Hartung, W., Al-Rasheid,
Khaled  A.S., Sonnewald, S., Sonnewald, U., Kneitz, S., Lachmann, N.,
Mendel, R.R., Bittner, F., Hetherington, A.M., and Hedrich, R. 2012. The
stomatal response to reduced relative humidity requires guard cell-
autonomous ABA synthesis. Current Biology 23, 53–57.

Stom ata a n d P ostharvest P hy s i o l o g y

Beerling, D.J., and Chaloner, W.G. 1993. The impact of atmospheric CO2 and
temperature-change on stomatal density—observations from quercus-
robur lammas leaves. Annals of Botany 71, 231–235.
Belin, C., De Franco, P.O., Bourbousse, C., Chaignepain, S., Schmitter, J.M.,
Vavasseur, A., Giraudat, J., Barbier-Brygoo, H., and Thomine, S. 2006.
Identification of features regulating OST1 kinase activity and OST1
function in guard cells. Plant Physiology 141, 1316–1327.
Ben-Yehoshua, S., and Rodov, V. 2003. Transpiration and water stress. In
Postharvest Physiology and Pathology of Vegetables, ed. J.A. Bartz and J.K.
Brecht, 111–159. 2nd ed. Marcel Dekker, New York.
Ben-Yehoshua, S., Kobiler, I., and Shapiro, B. 1979. Some physiological
effects of delaying deterioration of citrus fruits by individual seal pack-
aging in high density polyethylene film. Journal of the American Society
for Horticultural Science 104, 868–872.
Black, C.C., and Osmond, C.B. 2005. Crassulacean acid metabolism pho-
tosynthesis: ‘working the night shift’. In Discoveries in Photosynthesis,
ed. Govindjee, J.T. Beetty, H. Gest, and J.F. Allen, 881–893. Springer,
Dordrecht, The Netherlands.
Blanke, M. 1986. Comparative SEM study of stomata on developing quince,
apple, grape and tomato fruit. Angewandte Botanik 60, 209–214.
Blanke, M. 1987. Comparative SEM study of stomata and surface morphol-
ogy in apple. Angewandte Botanik 61, 433–438.
Blanke, M. 2002. Photosynthesis of strawberry fruit. Acta Horticulturae 567,
Blanke, M., and Lenz, F. 1989. Fruit photosynthesis. Plant, Cell and
Environment 12, 31–46.
Blanke, M.M. 1992. Photosynthesis of avocado fruit. In Proceedings of Second
World Avocado Congress, vol. 1, pp. 179–189. University of California,
Davis, CA.
Blanke, M.M. 1994. Regulation of respiration in apple, avocado and citrus
orange fruit. Acta Horticulturae 398, 139–146.
Blanke, M.M., and Leyhe, A. 1988. Stomatal and cuticular transpiration of the
cap and berry of grape. Journal of Plant Physiology 132, 250–253.
Blanke, M.M., Höfer, M., and Pring, R.J. 1994. Stomata and structure of tetra-
ploid apple leaves cultured in vitro. Annals of Botany 73, 651–654.
Blatt, M.R. 2000. Cellular signaling and volume control in stomatal movements
in plants. Annual Review of Cell and Developmental Biology, 16, 221–241.
Bohnert, H.J., Nelson, D.E., and Jensen, R.G. 1995. Adaptations to environ-
mental stresses. Plant Cell 7, 1099–1111.
Bradford, K.J. 1983. Involvement of plant growth substances in the altera-
tion of leaf gas exchange of flooded tomato plants. Plant Physiology 73,
Brainerd, K.E., and Fuchigami, L.H. 1982. Stomatal functioning of in vitro and
greenhouse apple leaves in darkness, mannitol, ABA, and CO2. Journal
of Experimental Botany 33, 388–392.


Bredmose, N.B., and Nielsen, K.L. 2009. Controlled atmosphere storage

at high CO2 and low O2 levels affects stomatal conductance and influ-
ence root formation in kalanchoe cuttings. Scientia Horticulturae 122,
Burchi, G., and Prisa, D. 2013. Preharvest conditions that can improve the
postharvest quality of ornamentals. Acta Horticulturae 970, 23–28.
Burg, S.P. 2004. Postharvest Physiology and Hypobaric Storage of Fresh Produce.
CABI Publishing, Wallingford, UK.
Cajuste, J.F., González-Candelas, L., Veyrat, A., García-Breijo, F.J., Reig-
Armiñana, J., and Lafuente, M.T. 2010. Epicuticular wax content
and morphology as related to ethylene and storage performance of
‘Navelate’ orange fruit. Postharvest Biology and Technology 55, 29–35.
Camejo, D., Rodríguez, P., Angeles Morales, M., Miguel Dell’Amico, J.,
Torrecillas, A., and Alarcón, J.J. 2005. High temperature effects on pho-
tosynthetic activity of two tomato cultivars with different heat suscepti-
bility. Journal of Plant Physiology 162, 281–289.
Camposeo, S., Palasciano, M., Vivaldi, G.A., and Godini, A. 2011. Effect of
increasing climatic water deficit on some leaf and stomatal parame-
ters of wild and cultivated almonds under Mediterranean conditions.
Scientia Horticulturae 127, 234–241.
Ceulemans, R., Heursel, J., Ibrahim, N., and Impens, I. 1984. Variations
among physiological, morphological and biochemical characteristics
of evergreen azalea (Rhododendron simsii Planch.) cultivars. Scientia
Horticulturae 22, 147–155.
Chen, L., Dodd, I.C., Davies, W.J., and Wilkinson, S. 2013. Ethylene limits
abscisic acid- or soil drying-induced stomatal closure in aged wheat
leaves. Plant, Cell and Environment 36, 1850–1859.
Chen, X., Pierik, R., Peeters, A.J.M., Poorter, H., Visser, E.J.W., Huber, H., de
Kroon, H., and Voesenek, L.A.C.J. 2010. Endogenous abscisic acid as a
key switch for natural variation in flooding-induced shoot elongation.
Plant Physiology 154, 969–977.
Cheng, W.H., Endo, A., Zhou, L., Penney, J., Chen, H.C., Arroyo, A., Leon, P.,
Nambara, E., Asami, T., and Seo, M. 2002. A unique short-chain dehy-
drogenase/reductase in Arabidopsis glucose signaling and abscisic acid
biosynthesis and functions. Plant Cell Online 14, 2723–2743.
Chimona, C., Stamellou, A., Argiropoulos, A., and Rhizopoulou, S. 2012. Study
of variegated and white flower petals of Capparis spinosa expanded at
dusk in arid landscapes. Journal of Arid Land 4, 171–179.
Christensen, J.V. 1976. Cracking in cherries. Danish Journal of Plant and Soil
Science 80, 289–324.
Christmann, A., Grill, E., and Huang, J. 2013. Hydraulic signals in long-dis-
tance signaling. Current Opinion in Plant Biology 16, 293–300.

Stom ata a n d P ostharvest P hy s i o l o g y

Christmann, A., Hoffmann, T., Teplova, I., Grill, E., and Muller, A. 2005.
Generation of active pools of abscisic acid revealed by in vivo imaging
of water-stressed arabidopsis. Plant Physiology 137, 209–219.
Christmann, A., Weiler, E.W., Steudle, E., and Grill, E. 2007. A hydraulic
signal in root-to-shoot signaling of water shortage. Plant Journal 52,
Coates, L., and Johnson, G. 1997. Postharvest diseases of fruit and vegeta-
bles. In Plant Pathogens and Plant Diseases, ed. J.F. Brown, and H.J.
Ogle, 533–548. Rockvale Publications, Armidale, Australia.
Cornish, K., King, A., Reid, M., and Paul, J. 1985. Role of ABA in stress-
induced reduction of water loss from potted chrysanthemum plants.
Acta Horticulturae 167, 381–386.
Crawford, A.J., McLachlan, D.H., Hetherington, A.M., and Franklin, K.A.
2012. High temperature exposure increases plant cooling capacity.
Current Biology 22, R396–R397.
Cutler, A.J., and Krochko, J.E. 1999. Formation and breakdown of ABA.
Trends in Plant Science 4, 472–478.
Cutler, S.R., Rodriguez, P.L., Finkelstein, R.R., and Abrams, S.R. 2010. Abscisic
acid: Emergence of a core signaling network. Annual Review of Plant
Biology 61, 651–679.
Daszkowska-Golec, A., and Szarejko, I. 2013. Open or close the gate—­
Stomata action under the control of phytohormones in drought stress
conditions. Frontiers in Plant Science 4, 138.
Davies, W., Kudoyarova, G., and Hartung, W. 2005. Long-distance ABA sig-
naling and its relation to other signaling pathways in the detection of
soil drying and the mediation of the plant’s response to drought. Journal
of Plant Growth Regulation 24, 285–295.
Desikan, R., Hancock, J.T., Bright, J., Harrison, J., Weir, I., Hooley, R., and
Neill, S.J. 2005. A role for ETR1 in hydrogen peroxide signaling in sto-
matal guard cells. Plant Physiology 137, 831–834.
Desikan, R., Last, K., Harrett-Williams, R., Tagliavia, C., Harter, K., Hooley,
R., Hancock, J.T., and Neill, S.J. 2006. Ethylene-induced stomatal clo-
sure in Arabidopsis occurs via AtrbohF-mediated hydrogen peroxide
synthesis. Plant Journal 47, 907–916.
Dietz, K.J., Sauter, A., Wichert, K., Messdaghi, D., and Hartung, W. 2000.
Extracellular β-glucosidase activity in barley involved in the hydrolysis
of ABA glucose conjugate in leaves. Journal of Experimental Botany 51,
Dodd, A.N., Salathia, N., Hall, A., Kévei, E., Tóth, R., Nagy, F., Hibberd, J.M.,
Millar, A.J., and Webb, A.A.R. 2005. Plant circadian clocks increase pho-
tosynthesis, growth, survival, and competitive advantage. Science 309,


Doheny-Adams, T., Hunt, L., Franks, P.J., Beerling, D.J., and Gray, J.E. 2012.
Genetic manipulation of stomatal density influences stomatal size, plant
growth and tolerance to restricted water supply across a growth car-
bon dioxide gradient. Philosophical Transactions of the Royal Society B:
Biological Sciences 367, 547–555.
Dong, X., Ramírez-Sánchez, M., Huber, D.J., Rao, J., Zhang, Z., Choi, S.T.,
and Lee, J.H. 2013. Diffusivity of 1-methylcyclopropene in spinach and
bok choi leaf tissue, disks of tomato and avocado fruit tissue, and whole
tomato fruit. Postharvest Biology and Technology 78, 40–47.
Drake, P.L., Froend, R.H., and Franks, P.J. 2013. Smaller, faster stomata: scal-
ing of stomatal size, rate of response, and stomatal conductance. Journal
of Experimental Botany 64, 495–505.
Dubovskaya, L.V., Bakakina, Y.S., Kolesneva, E.V., Sodel, D.L., McAinsh,
M.R., Hetherington, A.M., and Volotovski, I.D. 2011. cGMP-depen-
dent ABA-induced stomatal closure in the ABA-insensitive Arabidopsis
mutant abi1-1. New Phytologist 191, 57–69.
Effmert, U., Grosse, J., Rose, U.S.R., Ehrig, F., Kagi, R., and Piechulla, B.
2005. Volatile composition, emission pattern, and localization of floral
scent emission in Mirabilis jalapa (Nyctaginaceae). American Journal
of Botany 92, 2–12.
Elibox, W., and Umaharan, P. 2008. Morphophysiological characteristics
associated with vase life of cut flowers of anthurium. HortScience 43,
Elibox, W., and Umaharan, P. 2010. Cultivar differences in the deterioration
of vase-life in cut-flowers of Anthurium andraeanum is determined by
mechanisms that regulate water uptake. Scientia Horticulturae 124,
Elzenga, J.T.M., and Van Volkenburgh, E. 1997. Kinetics of Ca2+- and ATP-
dependent, voltage-controlled anion conductance in the plasma mem-
brane of mesophyll cells of Pisum sativum. Planta 201, 415–423.
Elzenga, J.T.M., Staal, M., and Prins, H.B.A. 2000. Modulation by phyto-
chrome of the blue light-induced extracellular acidification by leaf epi-
dermal cells of pea (Pisum sativum L.): a kinetic analysis. Plant Journal
22, 377–389.
Endo, A., Sawada, Y., Takahashi, H., Okamoto, M., Ikegami, K., Koiwai, H.,
Seo, M., Toyomasu, T., Mitsuhashi, W., Shinozaki, K., Nakazono, M.,
Kamiya, Y., Koshiba, T., and Nambara, T. 2008. Drought induction of
arabidopsis 9-cis-epoxycarotenoid dioxygenase occurs in vascular
parenchyma cells. Plant Physiology 147, 1984–1993.
Ezell, B.D., and Wilcox, M.S. 1959. Vegetable vitamins, loss of vitamin C
in fresh vegetables as related to wilting and temperature. Journal of
Agricultural and Food Chemistry 7, 507–509.
Ezell, B.B., and Wilcox, M.S. 1962. Vegetable vitamin values, loss of caro-
tene in fresh vegetables as related to wilting and temperature. Journal
of Agricultural and Food Chemistry 10, 124–126.

Stom ata a n d P ostharvest P hy s i o l o g y

Fallik, E., Grinberg, S., Alkalai, S., Yekutieli, O., Wiseblum, A., Regev, R.,
Beres, H., and Bar-Lev, E. 1999. A unique rapid hot water treatment
to improve storage quality of sweet pepper. Postharvest Biology and
Technology 15, 25–32.
Fanourakis, D., Carvalho, S.M.P., Almeida, D.P.F., and Heuvelink, E. 2011.
Avoiding high relative air humidity during critical stages of leaf ontogeny
is decisive for stomatal functioning. Physiologia Plantarum 142, 274–286.
Fanourakis, D., Heuvelink, E., and Carvalho, S.M.P. 2013a. A comprehensive
analysis of the physiological and anatomical components involved in
higher water loss rates after leaf development at high humidity. Journal
of Plant Physiology 170, 890–898.
Fanourakis, D., Pieruschka, R., Savvides, A., Macnish, A.J., Sarlikioti, V.,
and Woltering, E.J. 2013b. Sources of vase life variation in cut roses: a
review. Postharvest Biology and Technology 78, 1–15.
Feller, U. 2006. Stomatal opening at elevated temperature: an underestimated
regulatory mechanism? Genaral and Applied Plant Physiology 32, 19–31.
Ferrante, A., Vernieri, P., Serra, G., and Tognoni, F. 2004. Changes in abscisic
acid during leaf yellowing of cut stock flowers. Plant Growth Regulation
43, 127–134.
Fonseca, J.M., Kim, H.J., Kline, W.L., Wyenandt, C.A., Hoque, M., Ajwa,
H., and French, N. 2009. Effect of preharvest application of a second-
generation harpin protein on microbial quality, antioxidants, and shelf
life of fresh-cut lettuce. Journal of the American Society for Horticultural
Science 134, 141–147.
Fordham, M.C., Harrison-Murray, R.S., Knight, L., and Evered, C.E. 2001. Effects
of leaf wetting and high humidity on stomatal function in leafy cuttings and
intact plants of Corylus maxima. Physiologia Plantarum 113, 233–240.
Franck, C., Lammertyn, J., Ho, Q.T., Verboven, P., Verlinden, B., and Nicolaï,
B.M. 2007. Browning disorders in pear fruit. Postharvest Biology and
Technology 43, 1–13.
Franks, P.J., and Farquhar, G.D. 2007. The mechanical diversity of stomata
and its significance in gas-exchange control. Plant Physiology 143, 78–87.
Frechilla, S., Zhu, J., Talbott, L.D., and Zeiger, E. 1999. Stomata from npq1,
a zeaxanthin-less Arabidopsis mutant, lack a specific response to blue
light. Plant and Cell Physiology 40, 949–954.
Frommhold, I. 1982. Effect of ethephon on stomatal opening in detached
epidermal strips of tobacco leaves (Nicotiana tobacum L. cv. Samsun).
Biologia Plantarum 24, 303–306.
Fujii, H., and Zhu, J.K. 2009. Arabidopsis mutant deficient in 3 abscisic acid-
activated protein kinases reveals critical roles in growth, reproduction,
and stress. Proceedings of the National Academy of Sciences of the United
States of America 106, 8380–8385.
Fujii, H., Chinnusamy, V., Rodrigues, A., Rubio, S., Antoni, R., Park, S.Y.,
Cutler, S.R., Sheen, J., Rodriguez, P.L., and Zhu, J.K. 2009. In vitro recon-
stitution of an abscisic acid signaling pathway. Nature 462, 660–664.


Fujita, Y., Nakashima, K., Yoshida, T., Katagiri, T., Kidokoro, S., Kanamori,
N., Umezawa, T., Fujita, M., Maruyama, K., Ishiyama, K., Kobayashi,
M., Nakasone, S., Yamada, K., Ito, T., Shinozaki, K., and Yamaguchi-
Shinozaki, K. 2009. Three SnRK2 protein kinases are the main posi-
tive regulators of abscisic acid signaling in response to water stress in
Arabidopsis. Plant and Cell Physiology 50, 2123–2132.
Gan, S., and Amasino, R.M. 1995. Inhibition of leaf senescence by autoregu-
lated production of cytokinin. Science 270, 1986–1988.
Geiger, D., Scherzer, S., Mumm, P., Marten, I., Ache, P., Matschi, S., Liese, A.,
Wellmann, C., Al-Rasheid, K.A.S., Grill, E. et al. 2010. Guard cell anion
channel SLAC1 is regulated by CDPK protein kinases with distinct Ca2+
affinities. Proceedings of the National Academy of Sciences of the United
States of America 107, 8023–8028.
Geiger, D., Scherzer, S., Mumm, P., Stange, A., Marten, I., Bauer, H., Ache,
P., Matschi, S., Liese, A., Al-Rasheid, K.A.S., Grill, E., Romeis, T., and
Hedrich, R. 2009. Activity of guard cell anion channel SLAC1 is con-
trolled by drought-stress signaling kinase-phosphatase pair. Proceedings
of the National Academy of Sciences of the United States of America 106,
Ghashghaie, J., Brenckmann, F., and Saugier, B. 1992. Water relations and
growth of rose plants cultured in vitro under various relative humidities.
Plant Cell, Tissue and Organ Culture 30, 51–57.
Giday, H., Fanourakis, D., Kjaer, K.H., Fomsgaard, I.S., and Ottosen, C.O.
2013a. Foliar abscisic acid content underlies genotypic variation in sto-
matal responsiveness after growth at high relative air humidity. Annals
of Botany 112, 1857–1867.
Giday, H., Kjaer, K.H., Fanourakis, D., and Ottosen, C.O. 2013b. Smaller
stomata require less severe leaf drying to close: A case study in Rosa
hydrida. Journal of Plant Physiology 170, 1309–1316.
Giday, H., Fanourakis, D., Kjaer, K.H., Fomsgaard, I.S., and Ottosen, C.O.
2014. Threshold response of stomatal closing ability to leaf abscisic
acid concentration during growth. Journal of Experimental Botany, doi:
Gudesblat, G.E., Torres, P.S., and Vojnov, A.A. 2009. Stomata and pathogens:
Warfare at the gates. Plant Signaling and Behavior 4, 1114–1116.
Guo, Y., Xiong, L., Song, C.P., Gong, D., Halfter, U., and Zhu, J.K. 2002. A
calcium sensor and its interacting protein kinase are global regulators
of abscisic acid signaling in arabidopsis. Developmental Cell 3, 233–244.
Halevy, A.H., and Mayak, S. 1981. Senescence and postharvest physiology of
cut flowers. Part 2. Horticultural Reviews 3, 59–143.
Halevy, A.H., Mayak, S., Tirosh, T., Spiegelstein, H., and Kofranek, A.M.
1974. Opposing effects of abscisic acid on senescence of rose flowers.
Plant and Cell Physiology 15, 813–821.
Hall, A.E., Camacho, B.S.E., and Kaufmann, M.R. 1975. Regulation of water
loss by citrus leaves. Physiologia Plantarum 33, 62–65.

Stom ata a n d P ostharvest P hy s i o l o g y

Hao, J.H., Wang, X.L., Dong, C.J., Zhang, Z.G., and Shang, Q.M. 2011.
Salicylic acid induces stomatal closure by modulating endogenous hor-
mone levels in cucumber cotyledons. Russian Journal of Plant Physiology
58, 906–913.
Harada, A., and Shimazaki, K.I. 2009. Measurement of changes in cytosolic
Ca2+ in Arabidopsis guard cells and mesophyll cells in response to blue
light. Plant and Cell Physiology 50, 360–373.
Hauser, F., Waadt, R., and Schroeder, J.I. 2011. Evolution of abscisic acid syn-
thesis and signaling mechanisms. Current Biology 21, R346–R355.
Hazarika, B.N. 2006. Morpho-physiological disorders in in vitro culture of
plants. Scientia Horticulturae 108, 105–120.
Hentzen, A.E., Smart, L.B., Wimmers, L.E., Fang, H.H., Schroeder, J.I.,
and Bennett, A.B. 1996. Two plasma membrane H+ -ATPase genes
expressed in guard cells of Vicia faba are also expressed throughout
the plant. Plant and Cell Physiology 37, 650–659.
Herppich, W.B., Linke, M., Landahl, S., and Gzik, A. 2001. Preharvest and
postharvest responses of radish to reduced water supply during growth.
Acta Horticulturae 553, 89–90.
Hetherington, A.M., and Woodward, F.I. 2003. The role of stomata in sensing
and driving environmental change. Nature 424, 901–908.
Hew, C.S., Lee, G.L., and Wong, S.C. 1980. Occurrence of non-functional sto-
mata in the flowers of tropical orchids. Annals of Botany 46, 195–201.
Hewett, E.W. 2006. An overview of preharvest factors influencing postharvest
quality of horticultural products. International Journal of Postharvest
Technology and Innovation 1, 4–15.
Holbrook, N.M., Shashidhar, V., James, R.A., and Munns, R. 2002. Stomatal
control in tomato with ABA‐deficient roots: Response of grafted plants
to soil drying. Journal of Experimental Botany 53, 1503–1514.
Hossain, M.A., Munemasa, S., Uraji, M., Nakamura, Y., Mori, I.C., and Murata,
Y. 2011. Involvement of endogenous abscisic acid in methyl jasmonate-
induced stomatal closure in Arabidopsis. Plant Physiology 156, 430–438.
Hronková, M., Zahradníčková, H., Šimková, M., Šimek, P., and Heydová, A.
2003. The role of abscisic acid in acclimation of plants cultivated in vitro
to ex vitro conditions. Biologia Plantarum 46, 535–541.
Hu, X., Zhang, A., Zhang, J., and Jiang, M. 2006. Abscisic acid is a key inducer
of hydrogen peroxide production in leaves of maize plants exposed to
water stress. Plant and Cell Physiology 47, 1484–1495.
Huang, X.Y., Chao, D.Y., Gao, J.P., Zhu, M.Z., Shi, M., and Lin, H.X. 2009. A
previously unknown zinc finger protein, DST, regulates drought and salt
tolerance in rice via stomatal aperture control. Genes and Development
23, 1805–1817.
Hubbard, K.E., Nishimura, N., Hitomi, K., Getzoff, E.D., and Schroeder,
J.I. 2010. Early abscisic acid signal transduction mechanisms: Newly
discovered components and newly emerging questions. Genes and
Development 24, 1695–1708.


Hubbard, K.E., Siegel, R.S., Valerio, G., Brandt, B., and Schroeder, J.I. 2012.
Abscisic acid and CO2 signaling via calcium sensitivity priming in guard
cells, new CDPK mutant phenotypes and a method for improved resolu-
tion of stomatal stimulus–response analyses. Annals of Botany 109, 5–17.
Hunter, D.A., Ferrante, A., Vernieri, P., and Reid, M.S. 2004. Role of abscisic
acid in perianth senescence of daffodil (Narcissus pseudonarcissus
‘Dutch Master’). Physiologia Plantarum 121, 313–321.
In, B.C., Motomura, S., Inamoto, K., Doi, M., and Mori, G. 2007. Multivariate
analysis of relations between preharvest environmental factors, post-
harvest morphological and physiological factors, and vase life of cut
‘Asami Red’ roses. Journal of the Japanese Society for Horticultural
Science 76, 66–72.
Islam, N., Torre, S., Wold, A.B., and Gislerød, H.R. 2010. Effects of growing
conditions on the postharvest quality of herbs. Acta Horticulturae 877,
Iuchi, S., Kobayashi, M., Taji, T., Naramoto, M., Seki, M., Kato, T., Tabata,
S., Kakubari, Y., Yamaguchi‐Shinozaki, K., and Shinozaki, K. 2001.
Regulation of drought tolerance by gene manipulation of 9‐cis‐epoxy-
carotenoid dioxygenase, a key enzyme in abscisic acid biosynthesis in
Arabidopsis. Plant Journal 27, 325–333.
Jiang, F., and Hartung, W. 2008. Long-distance signaling of abscisic acid
(ABA): the factors regulating the intensity of the ABA signal. Journal of
Experimental Botany 59, 37–43.
Johnson, B.E., and Brun, W. 1966. Stomatal density and responsiveness of
banana fruit stomates. Plant Physiology 41, 99–101.
Jones, H.G. 1999. Use of thermography for quantitative studies of spatial and
temporal variation of stomatal conductance over leaf surfaces. Plant,
Cell and Environment 22, 1043–1055.
Joshi-Saha, A., Valon, C., and Leung, J. 2011. Abscisic acid signal off the
STARTing block. Molecular Plant 4, 562–580.
Khan, K., Joshi, P., and Purohit, S.D. 2010. Stomatal characteristics dur-
ing micropropagation of Wrightia tomentosa. Acta Horticulturae 865,
Khokon, M.A.R., Okuma, E., Hossain, M.A., Munemasa, S., Uraji, M.,
Nakamura, Y., Mori, I.C., and Murata, Y. 2011. Involvement of extra-
cellular oxidative burst in salicylic acid-induced stomatal closure in
Arabidopsis. Plant, Cell and Environment 34, 434–443.
Kim, T.H., Böhmer, M., Hu, H., Nishimura, N., and Schroeder, J.I. 2010.
Guard cell signal transduction network: advances in understanding
abscisic acid, CO2, and Ca2+ signaling. Annual Review of Plant Biology
61, 561–591.
Kirk, H., and Skytt Anderson, A. 1985. Influence of low pressure storage on sto-
matal opening and rooting of cuttings. Acta Horticulturae 181, 393–398.
Kirk, H., Andersen, A.S., Veierskov, B., Johansen, E., and Aabrandt, Z. 1986.
Low-pressure storage of hibiscus cuttings. Effect on stomatal opening
and rooting. Annals of Botany 58, 389–396.

Stom ata a n d P ostharvest P hy s i o l o g y

Koiwai, H., Nakaminami, K., Seo, M., Mitsuhashi, W., Toyomasu, T., and
Koshiba, T. 2004. Tissue-specific localization of an abscisic acid biosyn-
thetic enzyme, AAO3, in arabidopsis. Plant Physiology 134, 1697–1707.
Krochko, J.E., Abrams, G.D., Loewen, M.K., Abrams, S.R., and Cutler, A.J.
1998. (+)-Abscisic acid 8´-hydroxylase is a cytochrome P450 monooxy-
genase. Plant Physiology 118, 849–860.
Kroupitski, Y., Golberg, D., Belausov, E., Pinto, R., Swartzberg, D., Granot,
D., and Sela, S. 2009. Internalization of Salmonella enterica in leaves is
induced by light and involves chemotaxis and penetration through open
stomata. Applied and Environmental Microbiology 75, 6076–6086.
Kuromori, T., Sugimoto, E., and Shinozaki, K. 2014. Intertissue signal trans-
fer of abscisic acid from vascular cells to guard cells. Plant Physiology
164, 1587–1592.
Kushiro, T., Okamoto, M., Nakabayashi, K., Yamagishi, K., Kitamura, S.,
Asami, T., Hirai, N., Koshiba, T., Kamiya, Y., and Nambara, E. 2004. The
Arabidopsis cytochrome P450 CYP707A encodes ABA 8´-hydroxylases:
key enzymes in ABA catabolism. EMBO Journal 23, 1647–1656.
Kwak, J.M., Moon, J.H., Murata, Y., Kuchitsu, K., Leonhardt, N., DeLong, A.,
and Schroeder, J.I. 2002. Disruption of a guard cell–expressed protein
phosphatase 2A regulatory subunit, RCN1, confers abscisic acid insen-
sitivity in Arabidopsis. Plant Cell 14, 2849–2861.
Laurin, É., Nunes, M.C.N., Émond, J.P., and Brecht, J.K. 2006. Residual effect
of low-pressure stress during simulated air transport on Beit Alpha-type
cucumbers: Stomata behavior. Postharvest Biology and Technology 41,
Lazan, H., Ali, Z.M., Mohd, A.A., and Nahar, F. 1987a. Water stress and qual-
ity decline during storage of tropical leafy vegetables. Journal of Food
Science 52, 1286–1292.
Lazan, H., Ali, Z.M., Mohd, A.A., and Ong, G.B. 1987b. Influence of water
stress on cold-induced sweetening in leafy vegetable, Brassica juncea L.
Journal of Food Science 52, 1289–1292.
Lebaudy, A., Vavasseur, A., Hosy, E., Dreyer, I., Leonhardt, N., Thibaud, J.B.,
Véry, A.A., Simonneau, T., and Sentenac, H. 2008. Plant adaptation to
fluctuating environment and biomass production are strongly depen-
dent on guard cell potassium channels. Proceedings of the National
Academy of Sciences of the United States of America 105, 5271–5276.
Leckie, C.P., McAinsh, M.R., Montgomery, L., Priestley, A.J., Staxen, I.,
Webb, A.A.R., and Hetherington, A.M. 1998. Second messengers in
guard cells. Journal of Experimental Botany 49, 339–349.
Lee, K.H., Piao, H.L., Kim, H.Y., Choi, S.M., Jiang, F., Hartung, W., Hwang,
I., Kwak, J.M., and Lee, I.J. 2006. Activation of glucosidase via stress-
induced polymerization rapidly increases active pools of abscisic acid.
Cell 126, 1109–1120.
Lee, S.C., and Luan, S. 2012. ABA signal transduction at the crossroad of
biotic and abiotic stress responses. Plant, Cell and Environment 35,


Lee, S.C., Lan, W., Buchanan, B.B., and Luan, S. 2009. A protein kinase-phos-
phatase pair interacts with an ion channel to regulate ABA signaling in
plant guard cells. Proceedings of the National Academy of Sciences of the
United States of America 106, 21419–21424.
Lee, S.C., Lim, C.W., Lan, W., He, K., and Luan, S. 2013. ABA signaling in
guard cells entails a dynamic protein-protein interaction relay from the
PYL-RCAR family receptors to ion channels. Molecular Plant 6, 528–538.
Lee, S.K., and Kader, A.A. 2000. Preharvest and postharvest factors influenc-
ing vitamin C content of horticultural crops. Postharvest Biology and
Technology 20, 207–220.
Leonhardt, N., Vavasseur, A., and Forestier, C. 1999. ATP binding cassette
modulators control abscisic acid-regulated slow anion channels in
guard cells. Plant Cell 11, 1141–1151.
Leung, J., Merlot, S., and Giraudat, J. 1997. The Arabidopsis ABSCISIC ACID-
INSENSITIVE2 (ABI2) and ABI1 genes encode homologous protein
phosphatases 2C involved in abscisic acid signal transduction. Plant
Cell 9, 759–771.
Levchenko, V., Konrad, K.R., Dietrich, P., Roelfsema, M.R.G., and Hedrich, R.
2005. Cytosolic abscisic acid activates guard cell anion channels without
preceding Ca2+ signals. Proceedings of the National Academy of Sciences
of the United States of America 102, 4203–4208.
Li, J., and Assmann, S.M. 1996. An abscisic acid-activated and calcium-indepen-
dent protein kinase from guard cells of fava bean. Plant Cell 8, 2359–2368.
Li, S., Assmann, S.M., and Albert, R. 2006. Predicting essential components
of signal transduction networks: a dynamic model of guard cell abscisic
acid signaling. PLoS Biology 4, 1732–1748.
Li, X., Zhu, X., Zhao, N., Fu, D., Li, J., Chen, W., and Chen, W. 2013. Effects
of hot water treatment on anthracnose disease in papaya fruit and its
possible mechanism. Postharvest Biology and Technology 86, 437–446.
López-Gálvez, F., Gil, M.I., Truchado, P., Selma, M.V., and Allende, A. 2010.
Cross-contamination of fresh-cut lettuce after a short-term exposure
during pre-washing cannot be controlled after subsequent washing with
chlorine dioxide or sodium hypochlorite. Food Microbiology 27, 199–204.
Lu, Y.Y., and Chen, C.Y. 2005. Molecular analysis of lily leaves in response to
salicylic acid effective towards protection against Botrytis elliptica. Plant
Science 169, 1–9.
Ma, Y., Szostkiewicz, I., Korte, A., Moes, D., Yang, Y., Christmann, A., and
Grill, E. 2009. Regulators of PP2C phosphatase activity function as
abscisic acid sensors. Science 324, 1064–1068.
MacRobbie, E.A.C. 1990. Calcium-dependent and calcium-independent
events in the initiation of stomatal closure by abscisic acid. Proceedings
of the Royal Society B: Biological Sciences 241, 214–219.
Madhavan, S., Chrominiski, A., and Smith, B.N. 1983. Effect of ethylene
on stomatal opening in tomato and carnation leaves. Plant and Cell
Physiology 24, 569–572.

Stom ata a n d P ostharvest P hy s i o l o g y

Magwaza, L.S., Opara, U.L., Cronje, P.J.R., Landahl, S., and Terry, L.A. 2013.
Canopy position affects rind biochemical profile of ‘Nules Clementine’
mandarin fruit during postharvest storage. Postharvest Biology and
Technology 86, 300–308.
Marten, H., Hedrich, R., and Roelfsema, M.R.G. 2007a. Blue light inhibits
guard cell plasma membrane anion channels in a phototropin-depen-
dent manner. Plant Journal 50, 29–39.
Marten, H., Konrad, K.R., Dietrich, P., Roelfsema, M.R.G., and Hedrich, R.
2007b. Ca2+-dependent and -independent abscisic acid activation of
plasma membrane anion channels in guard cells of Nicotiana tabacum.
Plant Physiology 143, 28–37.
Martin, E.S., and Meidner, H. 1971. Endogenous stomatal movements in
Tradescantia virginiana. New Phytologist 70, 923–928.
Martínez-Sánchez, A., Tudela, J.A., Luna, C., Allende, A., and Gil, M.I. 2011.
Low oxygen levels and light exposure affect quality of fresh-cut Romaine
lettuce. Postharvest Biology and Technology 59, 34–42.
Mayak, S., and Dilley, D. 1976. Effect of sucrose on response of cut carnation
(Dianthus caryophyllus) to kinetin, ethylene, and abscisic acid. Journal
of the American Society for Horticultural Science 101, 583–585.
Mayak, S., and Halevy, A.H. 1974. The action of kinetin in improving the
water balance and delaying sensecence processes of cut rose flowers.
Physiologia Plantarum 32, 330–336.
Mayak, S., Halevy, A., and Katz, M. 1972. Correlative changes in phytohor-
mones in relation to senescence processes in rose petals. Physiologia
Plantarum 27, 1–4.
Melhorn, V., Matsumi, K., Koiwai, H., Ikegami, K., Okamoto, M., Nambara,
E., Bittner, F., and Koshiba, T. 2008. Transient expression of AtNCED3
and AAO3 genes in guard cells causes stomatal closure in Vicia faba.
Journal of Plant Research 121, 125–131.
Melotto, M., Underwood, W., Koczan, J., Nomura, K., and He, S.Y. 2006. Plant
stomata function in innate immunity against bacterial invasion. Cell 126,
Melotto, M., Underwood, W., and Sheng, Y.H. 2008. Role of stomata in
plant innate immunity and foliar bacterial diseases. Annual Review of
Phytopathology 46, 101–122.
Mencarelli, F., and Bellincontro, A. 2013. Technology and management of
postharvest dehydration. In Sweet, Reinforced and Fortified Wines:
Grape Biochemistry, Technology and Vinification, ed. F. Mencarelli and
P. Tonutti, 51–75. John Wiley & Son, Chichester, West Sussex, UK.
Mendgen, K., Hahn, M., and Deising, H. 1996. Morphogenesis and mech-
anisms of penetration by plant pathogenic fungi. Annual Review of
Phytopathology 34, 367–386.
Mengel, K., and Kirkby, E.A. 1982. Principles of Plant Nutrition, 655.
International Potash Institute, Bern.


Merlot, S., Leonhardt, N., Fenzi, F., Valon, C., Costa, M., Piette, L., Vavasseur,
A., Genty, B., Boivin, K., Muller, A., Giraudat, J., and Leung, J. 2007.
Constitutive activation of a plasma membrane H+ -ATPase prevents
abscisic acid-mediated stomatal closure. EMBO Journal 26, 3216–3226.
Merlot, S., Mustilli, A.C., Genty, B., North, H., Lefebvre, V., Sotta, B.,
Vavasseur, A., and Giraudat, J. 2002. Use of infrared thermal imaging
to isolate Arabidopsis mutants defective in stomatal regulation. Plant
Journal 30, 601–609.
Merritt, F., Kemper, A., and Tallman, G. 2001. Inhibitors of ethylene synthe-
sis inhibit auxin-induced stomatal opening in epidermis detached from
leaves of Vicia faba L. Plant and Cell Physiology 42, 223–230.
Milillo, S.R., Badamo, J.M., Boor, K.J., and Wiedmann, M. 2008. Growth
and persistence of Listeria monocytogenes isolates on the plant model
Arabidopsis thaliana. Food Microbiology 25, 698–704.
Mintz-Oron, S., Mandel, T., Rogachev, I., Feldberg, L., Lotan, O., Yativ, M.,
Wang, Z., Jetter, R., Venger, I., and Adato, A. 2008. Gene expression
and metabolism in tomato fruit surface tissues. Plant Physiology 147,
Moes, D., Himmelbach, A., Korte, A., Haberer, G., and Grill, E. 2008. Nuclear
localization of the mutant protein phosphatase abi1 is required for insen-
sitivity towards ABA responses in Arabidopsis. Plant Journal 54, 806–819.
Montillet, J.L., Leonhardt, N., Mondy, S., Tranchimand, S., Rumeau, D.,
Boudsocq, M., Garcia, A.V., Douki, T., Bigeard, J., Laurière, C.,
Chevalier, A., Castresana, C., and Hirt, H. 2013. An abscisic acid-
independent oxylipin pathway controls stomatal closure and immune
defense in Arabidopsis. PLoS Biology 11, e1001513. doi: 10.1371/journal.
Moran, R.E., DeEll, J.R., and Halteman, W. 2009. Effects of preharvest pre-
cipitation, air temperature, and humidity on the occurrence of soft scald
in ‘Honeycrisp’ apples. HortScience 44, 1645–1647.
Moreshet, S., and Green, G. 1980. Photosynthesis and diffusion conduc-
tance of the Valencia orange fruit under field conditions. Journal of
Experimental Botany 31, 15–27.
Mori, I.C., Murata, Y., Yang, Y., Munemasa, S., Wang, Y.F., Andreoli, S., Tiriac,
H., Alonso, J.M., Harper, J.F., Ecker, J.R., Kwak, J.R., and Schroeder, J.I.
2006. CDPKs CPK6 and CPK3 function in ABA regulation of guard cell
S-type anion- and Ca2+-permeable channels and stomatal closure. PLoS
Biology 4, 1749–1762.
Morison, J.I., and Gifford, R.M. 1983. Stomatal sensitivity to carbon dioxide
and humidity: A comparison of two C3 and two C4 grass species. Plant
Physiology 71, 789–796.
Mortensen, L.M., and Fjeld, T. 1998. Effects of air humidity, lighting period
and lamp type on growth and vase life of roses. Scientia Horticulturae
73, 229–237.

Stom ata a n d P ostharvest P hy s i o l o g y

Mortensen, L.M., and Gislerød, H.R. 1999. Influence of air humidity and light-
ing period on growth, vase life and water relations of 14 rose cultivars.
Scientia Horticulturae 82, 289–298.
Mortensen, L.M., and Gislerod, H.R. 2000. Effect of air humidity on growth,
keeping quality, water relations, and nutrient content of cut roses.
Gartenbauwissenschaft 65, 40–44.
Mortensen, L.M., and Gislerød, H.R. 2011. Vase life: the influence of variation
in air humidity, temperature and super-elevated CO2 concentration in
roses grown under continuous light. European Journal of Horticultural
Science 76, 63–68.
Mulas, M., and Schirra, M. 2007. The effect of heat conditioning treatments
on the postharvest quality of horticultural crops. Stewart Postharvest
Review 3.
Murata, Y., and Mori, I.C. 2013. Stomatal regulation of plant water status. In
Plant Abiotic Stress, ed. M.A. Jenks and P.M. Hasegawa, 47–67. John
Wiley & Son, West Sussex, UK.
Nambara, E., and Marion-Poll, A. 2005. Abscisic acid biosynthesis and catabo-
lism. Annual Review of Plant Biology 56, 165–185.
Negi, J., Matsuda, O., Nagasawa, T., Oba, Y., Takahashi, H., Kawai-Yamada,
M., Uchimiya, H., Hashimoto, M., and Iba, K. 2008. CO2 regulator
SLAC1 and its homologues are essential for anion homeostasis in plant
cells. Nature 452, 483–486.
Neill, S., Barros, R., Bright, J., Desikan, R., Hancock, J., Harrison, J., Morris,
P., Ribeiro, D., and Wilson, I. 2008. Nitric oxide, stomatal closure, and
abiotic stress. Journal of Experimental Botany 59, 165–176.
Njombolwana, N.S., Erasmus, A., van Zyl, J.G., du Plooy, W., Cronje, P.J.R.,
and Fourie, P.H. 2013. Effects of citrus wax coating and brush type on
imazalil residue loading, green mould control and fruit quality retention
of sweet oranges. Postharvest Biology and Technology 86, 362–371.
Nobel, P.S. 1999. Physicochemical and Environmental Plant Physiology. 2nd
ed. Academic Press, San Diego.
Noichinda, S., Bodhipadma, K., Mahamontri, C., Narongruk, T., and Ketsa, S.
2007. Light during storage prevents loss of ascorbic acid, and increases
glucose and fructose levels in Chinese kale Brassica oleracea var. albo-
glabra. Postharvest Biology and Technology 44, 312–315.
Novák, V., and Vidovič, J. 2003. Transpiration and nutrient uptake dynamics
in maize (Zea mays L.). Ecological Modelling 166, 99–107.
Nunes, M.C.N., Brecht, J.K., Morais, A.M.M.B., and Sargent, S.A. 1998.
Controlling temperature and water loss to maintain ascorbic acid levels
in strawberries during postharvest handling. Journal of Food Science 63,
Oh, M.M., Trick, H.N., and Rajashekar, C.B. 2009. Secondary metabolism
and antioxidants are involved in environmental adaptation and stress
tolerance in lettuce. Journal of Plant Physiology 166, 180–191.


Okamoto, M., Tanaka, Y., Abrams, S.R., Kamiya, Y., Seki, M., and Nambara,
E. 2009. High humidity induces abscisic acid 8´-hydroxylase in stomata
and vasculature to regulate local and systemic abscisic acid responses
in Arabidopsis. Plant Physiology 149, 825–834.
Orsini, F., Alnayef, M., Bona, S., Maggio, A., and Gianquinto, G. 2012. Low
stomatal density and reduced transpiration facilitate strawberry adapta-
tion to salinity. Environmental and Experimental Botany 81, 1–10.
Osakabe, Y., Yamaguchi-Shinozaki, K., Shinozaki, K., and Tran, L.S.P. 2013.
ABA control of plant macroelement membrane transport systems in
response to water deficit and high salinity. New Phytologist 202, 35–49.
Outlaw, W.H. 2003. Integration of cellular and physiological functions of
guard cells. Critical Reviews in Plant Sciences 22, 503–529.
Pallaghy, C., and Raschke, K. 1972. No stomatal response to ethylene. Plant
Physiology 49, 275–276.
Pallas, J.E., and Kays, S.J. 1982. Inhibition of photosynthesis by ethylene—
a stomatal effect. Plant Physiology 70, 598–601.
Pandey, R., Chacko, P.M., Choudhary, M.L., Prasad, K.V., and Pal, M.
2007. Higher than optimum temperature under CO2 enrichment influ-
ences stomata anatomical characters in rose (Rosa hybrida). Scientia
Horticulturae 113, 74–81.
Pandey, R., Chacko, P.M., Pal, M., Choudhary, M.L., and Singh, R. 2010.
Influence of growth temperature on keeping quality traits of rose (Rosa
hybrida L.) cut flowers grown under continuous CO2 enrichment. Indian
Journal of Plant Physiology 15, 343–349.
Parés, J., Arizaleta, M., Sanabria, M.E., and García, G. 2008. Effect of salin-
ity levels on the stomatal density, stomatal index and leaf thickness of
Carica papaya L. Acta Botánica Venezuelica 31, 27–34.
Park, S.Y., Fung, P., Nishimura, N., Jensen, D.R., Fujii, H., Zhao, Y., Lumba,
S., Santiago, J., Rodrigues, A., Chow, T.F.F., Alfred, S.E., Bonetta, D.,
Finkelstein, R., Provart, N.J., Desveaux, D., Rodriguez, P.L., McCourt,
P., Zhu, J.K., Schroeder, J.I., Volkman, B.F., and Cutler, S.R. 2009.
Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL
family of START proteins. Science 324, 1068–1071.
Parvathi, K., and Raghavendra, A.S. 1997. Blue light-promoted stomatal open-
ing in abaxial epidermis of Commelina benghalensis is maximal at low
calcium. Physiologia Plantarum 101, 861–864.
Paull, R.E. 1982. Anthurium (Anthurium andraeanum Andre) vase life evalu-
ation criteria. HortScience 17, 606–607.
Paull, R.E. 1999. Effect of temperature and relative humidity on fresh com-
modity quality. Postharvest Biology and Technology 15, 263–277.
Peana, A.T., D’Aquila, P.S., Panin, F., Serra, G., Pippia, P., and Moretti, M.D.L.
2002. Anti-inflammatory activity of linalool and linalyl acetate constitu-
ents of essential oils. Phytomedicine 9, 721–726.

Stom ata a n d P ostharvest P hy s i o l o g y

Peschel, S., Beyer, M., and Knoche, M. 2003. Surface characteristics of sweet
cherry fruit: stomata-number, distribution, functionality and surface
wetting. Scientia Horticulturae 97, 265–278.
Peterson, K.M., Rychel, A.L., and Torii, K.U. 2010. Out of the mouths of
plants: The molecular basis of the evolution and diversity of stomatal
development. Plant Cell 22, 296–306.
Pettersen, R.I., Moe, R., and Gislerød, H.R. 2007. Growth of pot roses and post-
harvest rate of water loss as affected by air humidity and temperature
variations during growth under continuous light. Scientia Horticulturae
114, 207–213.
Pettersen, R.I., Mortensen, L.M., Moe, R., and Gislerød, H.R. 2006. Air humid-
ity control essential for rose production under continuous lighting. Acta
Horticulturae 711, 323–331.
Pompodakis, N.E., Joyce, D.C., Terr y, L.A., and L ydakis, D.E. 2004.
Effects of vase solution pH and abscisic acid on the longevity of cut
‘Baccara’ roses. Journal of Horticultural Science and Biotechnology
79, 828–832.
Porat, R., Daus, A., Weiss, B., Cohen, L., Fallik, E., and Droby, S. 2000.
Reduction of postharvest decay in organic citrus fruit by a short hot
water brushing treatment. Postharvest Biology and Technology 18,
Pospíšilová, J., Synková, H., and Rulcová, J. 2000. Cytokinins and water stress.
Biologia Plantarum 43, 321–328.
Priest, D.M., Ambrose, S.J., Vaistij, F.E., Elias, L., Higgins, G.S., Ross, A.R.,
Abrams, S.R., and Bowles, D.J. 2006. Use of the glucosyltransferase
UGT71B6 to disturb abscisic acid homeostasis in Arabidopsis thaliana.
Plant Journal 46, 492–502.
Qin, X., and Zeevaart, J.A. 1999. The 9-cis-epoxycarotenoid cleavage reaction
is the key regulatory step of abscisic acid biosynthesis in water-stressed
bean. Proceedings of the National Academy of Sciences of the United States
of America 96, 15354–15361.
Qin, X., and Zeevaart, J.A.D. 2002. Overexpression of a 9-cis-epoxycarotenoid
dioxygenase gene in Nicotiana plumbaginifolia increases abscisic
acid and phaseic acid levels and enhances drought tolerance. Plant
Physiology 128, 544–551.
Raghavendra, A.S., Gonugunta, V.K., Christmann, A., and Grill, E. 2010. ABA
perception and signaling. Trends in Plant Science 15, 395–401.
Reid, M.S., and Jiang, C.Z. 2012. Postharvest biology and technology of cut
flowers and potted plants. Horticultural Reviews 40, 1–54.
Reynolds-Henne, C.E., Langenegger, A., Mani, J., Schenk, N., Zumsteg, A.,
and Feller, U. 2010. Interactions between temperature, drought and sto-
matal opening in legumes. Environmental and Experimental Botany 68,


Rezaei Nejad, A., and van Meeteren, U. 2005. Stomatal response character-
istics of Tradescantia virginiana grown at high relative air humidity.
Physiologia Plantarum 125, 324–332.
Rezaei Nejad, A., and van Meeteren, U. 2007. The role of abscisic acid in
disturbed stomatal response characteristics of Tradescantia virginiana
during growth at high relative air humidity. Journal of Experimental
Botany 58, 627–636.
Rezaei Nejad, A., and van Meeteren, U. 2008. Dynamics of adaptation of stoma-
tal behaviour to moderate or high relative air humidity in Tradescantia
virginiana. Journal of Experimental Botany 59, 289–301.
Rezaei Nejad, A., Harbinson, J., and van Meeteren, U. 2006. Dynamics of spa-
tial heterogeneity of stomatal closure in Tradescantia virginiana altered
by growth at high relative air humidity. Journal of Experimental Botany
57, 3669–3678.
Riederer, M., and Muller, C. 2008. Annual Plant Reviews, Biology of the Plant
Cuticle 23. John Wiley & Son, West Sussex, UK.
Rogers, C.A., Powell, R.D., and Sharpe, P.J.H. 1979. Relationship of tempera-
ture to stomatal aperture and potassium accumulation in guard cells of
Vicia faba. Plant Physiology 63, 388–391.
Rogiers, S.Y., Hatfield, J.M., Jaudzems, V.G., White, R.G., and Keller, M.
2004. Grape berry cv. Shiraz epicuticular wax and transpiration during
ripening and preharvest weight loss. American Journal of Enology and
Viticulture 55, 121–127.
Romero, P., Rodrigo, M.J., and Lafuente, M.T. 2013. Differential expression
of the Citrus sinensis ABA perception system genes during postharvest
fruit dehydration. Postharvest Biology and Technology 76, 65–73.
Romero-Aranda, R., Cantó-Garay, R., and Martínez, P.F. 1994. Distribution
and density of stomata in two cultivars of Gerbera jamesonii and its rela-
tion to leaf conductance. Scientia Horticulturae 58, 167–173.
Roychoudhury, A., Paul, S., and Basu, S. 2013. Cross-talk between abscisic
acid-dependent and abscisic acid-independent pathways during abiotic
stress. Plant Cell Reports 32, 985–1006.
Runkle, E., Woolard, D., Campbell, C., Blanchard, M., and Newton, L. 2007.
Exogenous applications of abscisic acid improved the postharvest
drought tolerance of several annual bedding plants. Acta Horticulturae
755, 127–133.
Rutland, R.B., Chang, H.L.L., and Pallas, J.E., Jr. 1987. Stomatal density of
snapdragon as a possible determinant of transpiration. HortScience 22,
Sachs, T. 1991. Stomata as an example of meristmoid development. In Pattern
Formation in Plant Tissues, 101–117. Cambridge University Press,
Sack, F.D. 1987. The development and structure of stomata. In Stomatal
Function, ed. E. Zeiger, G.D. Farquhar, and I.R. Cowan, 59–89. Stanford
University Press, Stanford, CA.

Stom ata a n d P ostharvest P hy s i o l o g y

Saito, S., Hirai, N., Matsumoto, C., Ohigashi, H., Ohta, D., Sakata, K., and
Mizutani, M. 2004. Arabidopsis CYP707As encode (+)-abscisic acid
8´-hydroxylase, a key enzyme in the oxidative catabolism of abscisic
acid. Plant Physiology 134, 1439–1449.
Sams, C.E. 1999. Preharvest factors affecting postharvest texture. Postharvest
Biology and Technology 15, 249–254.
Santamaria, J.M., and Kerstiens, G. 1994. The lack of control of water loss
in micropropagated plants is not related to poor cuticle development.
Physiologia Plantarum 91, 191–195.
Santamaria, J.M., Davies, W.J., and Atkinson, C.J. 1993. Stomata of microprop-
agated Delphinium plants respond to ABA, CO2, light and water poten-
tial, but fail to close fully. Journal of Experimental Botany 44, 99–107.
Santiago, J., Rodrigues, A., Saez, A., Rubio, S., Antoni, R., Dupeux, F., Park,
S.Y., Márquez, J.A., Cutler, S.R., and Rodriguez, P.L. 2009. Modulation
of drought resistance by the abscisic acid receptor PYL5 through inhibi-
tion of clade A PP2Cs. Plant Journal 60, 575–588.
Sauter, A., Dietz, K.J., and Hartung, W. 2002. A possible stress physiological
role of abscisic acid conjugates in root to shoot signaling. Plant, Cell and
Environment 25, 223–228.
Savvides, A., Fanourakis, D., and van Ieperen, W. 2012. Co-ordination of
hydraulic and stomatal conductances across light qualities in cucumber
leaves. Journal of Experimental Botany 63, 1135–1143.
Schirra, M., D’Hallewin, G., Ben-Yehoshua, S., and Fallik, E. 2000. Host–
pathogen interactions modulated by heat treatment. Postharvest Biology
and Technology 21, 71–85.
Schreiber, U., and Berry, J.A. 1977. Heat-induced changes of chlorophyll fluo-
rescence in intact leaves correlated with damage of the photosynthetic
apparatus. Planta 136, 233–238.
Schroeder, J.I., and Keller, B.U. 1992. Two types of anion channel cur-
rents in guard cells with distinct voltage regulation. Proceedings of
the National Academy of Sciences of the United States of America 89,
Schroeder, K.R., and Stimart, D.P. 2005. Comparison of stomatal density
and postharvest transpiration between long- and short-lived cut flower
genotypes of Antirrhinum majus L. Journal of the American Society for
Horticultural Science 130, 742–746.
Schroeder, J.I., Allen, G.J., Hugouvieux, V., Kwak, J.M., and Waner, D. 2001.
Guard cell signal transduction. Annual Review of Plant Biology, 52,
Schulze, E.D., Lange, O.L., Kappen, L., Buschbom, U., and Evenari, M. 1973.
Stomatal responses to changes in temperature at increasing water
stress. Planta 110, 29–42.
Schwertfeger, G., and Buchloh, G. 1968. Die Korkentwicklung in den
Abschlussgeweben der Triebe und Fruchte verschiedener Apfelsorten.
Gartenbauwissenschaft 33, 77–102.


Shamaila, M. 2005. Water and its relation to fresh produce. In Produce

Degradation: Pathways and Prevention, ed. S.I. Dike Ukuku and
O. Lamikanra, 267–291. CRC Press, Boca Raton, FL.
She, X., and Song, X. 2012. Ethylene inhibits abscisic acid-induced stomatal
closure in Vicia faba via reducing nitric oxide levels in guard cells. New
Zealand Journal of Botany 50, 203–216.
Shimizu-Yumoto, H., and Ichimura, K. 2009. Abscisic acid, in combination with
sucrose, is effective as a pulse treatment to suppress leaf damage and
extend foliage vase-life in cut Eustoma flowers. Journal of Horticultural
Science and Biotechnology 84, 107–111.
Shtein, I., Meir, S., Riov, J., and Philosoph-Hadas, S. 2011. Interconnection
of seasonal temperature, vascular traits, leaf anatomy and hydraulic
performance in cut Dodonaea ‘Dana’ branches. Postharvest Biology and
Technology 61, 184–192.
Siegel, R.S., Xue, S., Murata, Y., Yang, Y., Nishimura, N., Wang, A., and
Schroeder, J.I. 2009. Calcium elevation-dependent and attenuated rest-
ing calcium-dependent abscisic acid induction of stomatal closure and
abscisic acid-induced enhancement of calcium sensitivities of S-type
anion and inward-rectifying K+ channels in Arabidopsis guard cells.
Plant Journal 59, 207–220.
Sirichandra, C., Wasilewska, A., Vlad, F., Valon, C., and Leung, J. 2009. The
guard cell as a single-cell model towards understanding drought tolerance
and abscisic acid action. Journal of Experimental Botany 60, 1439–1463.
Sisler, E.C., and Wood, C. 1988. Interaction of ethylene and CO2. Physiologia
Plantarum 73, 440–444.
Slootweg, G., and van Meeteren, U. 1991. Transpiration and stomatal con-
ductance of roses cv. Sonia grown with supplemental lighting. Acta
Horticulturae, 298, 119–125.
Smith, D.L., Stommel, J.R., Fung, R.W.M., Wang, C.Y., and Whitaker, B.D.
2006. Influence of cultivar and harvest method on postharvest storage
quality of pepper (Capsicum annuum L.) fruit. Postharvest Biology and
Technology 42, 243–247.
Smith, K., Fleming, H., Van Dyke, C., and Lower, R. 1979. Scanning electron
microscopy of the surface of the pickling cucumber fruit. Journal of the
American Society for Horticultural Science 104, 528–533.
Song, C.P., Agarwal, M., Ohta, M., Guo, Y., Halfter, U., Wang, P., and Zhu, J.K.
2005. Role of an Arabidopsis AP2/EREBP-type transcriptional repressor
in abscisic acid and drought stress responses. Plant Cell 17, 2384–2396.
Song, X.G., She, X.P., and Wang, J. 2012. Inhibition of darkness-induced sto-
matal closure by ethylene involves a removal of hydrogen peroxide
from guard cells of Vicia faba. Russian Journal of Plant Physiology 59,
Song, X.G., She, X.P., He, J.M., Huang, C., and Song, T.S. 2006. Cytokinin- and
auxin-induced stomatal opening involves a decrease in levels of hydro-
gen peroxide in guard cells of Vicia faba. Functional Plant Biology 33,

Stom ata a n d P ostharvest P hy s i o l o g y

Sreenivasulu, N., Harshavardhan, V.T., Govind, G., Seiler, C., and Kohli, A.
2012. Contrapuntal role of ABA: Does it mediate stress tolerance or
plant growth retardation under long-term drought stress? Gene 506,
Stael, S., Wurzinger, B., Mair, A., Mehlmer, N., Vothknecht, U.C., and Teige,
M. 2011. Plant organellar calcium signaling: An emerging field. Journal
of Experimental Botany 63, 1525–1542.
Su, J.S., Wang, Y.F., Frelet, A., Leonhardt, N., Klein, M., Forestier, C., Mueller-
Roeber, B., Cho, M.H., Martinoia, E., and Schroeder, J.I. 2007. The
ATP binding cassette transporter AtMRP5 modulates anion and cal-
cium channel activities in Arabidopsis guard cells. Journal of Biological
Chemistry 282, 1916–1924.
Suhita, D., Raghavendra, A.S., Kwak, J.M., and Vavasseur, A. 2004.
Cytoplasmic alkalization precedes reactive oxygen species production
during methyl jasmonate- and abscisic acid-induced stomatal closure.
Plant Physiology 134, 1536–1545.
Sutter, J.U., Sieben, C., Hartel, A., Eisenach, C., Thiel, G., and Blatt, M.R.
2007. Abscisic acid triggers the endocytosis of the Arabidopsis KAT1 K+
channel and its recycling to the plasma membrane. Current Biology 17,
Swanson, B.T., Jr., Wilkins, H.F., Weiser, C.F., and Klein, I. 1975. Endogenous
ethylene and abscisic acid relative to phytogerontology. Plant Physiology
55, 370–376.
Tahir, S.S., and Rajput, M.T.M. 2010. SEM studies of petal structure of corolla
of the species Sibbaldia L. (Rosaceae). Pakistan Journal of Botany 42,
Takemiya, A., Kinoshita, T., Asanuma, M., and Shimazaki, K.I. 2006. Protein
phosphatase 1 positively regulates stomatal opening in response to blue
light in Vicia faba. Proceedings of the National Academy of Sciences of the
United States of America 103, 13549–13554.
Talbott, L., and Zeiger, E. 1998. The role of sucrose in guard cell osmoregula-
tion. Journal of Experimental Botany 49, 329–337.
Tallman, G. 2004. Are diurnal patterns of stomatal movement the result of
alternating metabolism of endogenous guard cell ABA and accumula-
tion of ABA delivered to the apoplast around guard cells by transpira-
tion? Journal of Experimental Botany 55, 1963–1976.
Tan, B.C., Joseph, L.M., Deng, W.T., Liu, L., Li, Q.B., Cline, K., and McCarty,
D.R. 2003. Molecular characterization of the Arabidopsis 9-cis-epoxyca-
rotenoid dioxygenase gene family. Plant Journal 35, 44–56.
Tanaka, Y., Sano, T., Tamaoki, M., Nakajima, N., Kondo, N., and Hasezawa,
S. 2005. Ethylene inhibits abscisic acid-induced stomatal closure in
Arabidopsis. Plant Physiology 138, 2337–2343.
Tanaka, Y., Sano, T., Tamaoki, M., Nakajima, N., Kondo, N., and Hasezawa,
S. 2006. Cytokinin and auxin inhibit abscisic acid-induced stomatal
closure by enhancing ethylene production in Arabidopsis. Journal of
Experimental Botany 57, 2259–2266.


Tang, J., Mitcham, E., Wang, S., and Lurie, S. 2007. Heat Treatments for
Postharvest Pest Control: Theory and Practice. CABI, Wallingford, UK.
Taylor, G.E., and Gunderson, C.A. 1986. The response of foliar gas exchange
to exogenously applied ethylene. Plant Physiology 82, 653–657.
Taylor, G.E., and Gunderson, C.A. 1988. Physiological site of ethylene effects
on carbon dioxide assimilation in Glycine max L. Merr. Plant Physiology
86, 85–92.
Tesfay, S.Z., Bertling, I., and Bower, J.P. 2011. Effects of postharvest potassium
silicate application on phenolics and other anti-oxidant systems aligned to
avocado fruit quality. Postharvest Biology and Technology 60, 92–99.
Thimann, K.V. 1985. The interaction of hormonal and environmental factors
in leaf senescence. Biologia Plantarum 27, 83–91.
Thimann, K.V., and Satler, S. 1979. Relation between senescence and stomatal
opening: senescence in darkness. Proceedings of the National Academy
of Sciences of the United States of America 76, 2770–2773.
Thomson, G.E. 2005. Postharvest handling effects on stomatal aperture of
harvested Brassica rapa var. perviridis ‘Komatsuna’ and B. juncea ‘Red
Giant’ mustard leaves. New Zealand Journal of Crop and Horticultural
Science 33, 311–316.
Tibaldi, G., Fontana, E., and Nicola, S. 2011. Growing conditions and posthar-
vest management can affect the essential oil of Origanum vulgare L. ssp.
hirtum (Link) Ietswaart. Industrial Crops and Products 34, 1516–1522.
Tijskens, L.M.M., Veltman, R.H., Heuvelink, E., and Simčič, M. 2003.
Modelling postharvest quality behaviour as affected by preharvest con-
ditions. Acta Horticulturae 599, 469–477.
Tissera, P., and Ayres, P. 1986. Endogenous ethylene affects the behaviour of
stomata in epidermis isolated from rust infected faba bean (Vicia faba
L.). New Phytologist 104, 53–61.
Torre, S. 1999. The role of air humidity and calcium in determining posthar-
vest quality of roses. Institutt for Plantefag, Norges Landbrukshogskole
[Department of Horticulture and Crop Sciences, Agricultural University
of Norway].
Torre, S., and Fjeld, T. 2001. Water loss and postharvest characteristics of
cut roses grown at high or moderate relative air humidity. Scientia
Horticulturae 89, 217–226.
Torre, S., Fjeld, T., and Gislerød, H.R. 2001. Effects of air humidity and K/Ca
ratio in the nutrient supply on growth and postharvest characteristics of
cut roses. Scientia Horticulturae 90, 291–304.
Torre, S., Fjeld, T., Gislerød, H.R., and Moe, R. 2003. Leaf anatomy and sto-
matal morphology of greenhouse roses grown at moderate or high air
humidity. Journal of the American Society for Horticultural Science 128,
Tricker, P.J., George Gibbings, J., Rodríguez López, C.M., Hadley, P., and
Wilkinson, M.J. 2012. Low relative humidity triggers RNA-directed de
novo DNA methylation and suppression of genes controlling stomatal
development. Journal of Experimental Botany 63, 3799–3814.

Stom ata a n d P ostharvest P hy s i o l o g y

Trontin, C., Tisné, S., Bach, L., and Loudet, O. 2011. What does Arabidopsis
natural variation teach us (and does not teach us) about adaptation in
plants? Current Opinion in Plant Biology 14, 225–231.
Tudela, J.A., Marín, A., Martínez-Sánchez, A., Luna, M.C., and Gil, M.I. 2013.
Preharvest and postharvest factors related to off-odours of fresh-cut ice-
berg lettuce. Postharvest Biology and Technology 86, 463–471.
Turrell, F., and Klotz, L. 1940. Density of stomata and oil glands and incidence
of water spot in the rind of Washington navel orange. Botanical Gazette
101, 862–871.
Tyree, M.T., and Dixon, M.A. 1986. Water stress induced cavitation and
embolism in some woody plants. Physiologia Plantarum 66, 397–405.
Umezawa, T., Sugiyama, N., Mizoguchi, M., Hayashi, S., Myouga, F.,
Yamaguchi-Shinozaki, K., Ishihama, Y., Hirayama, T., and Shinozaki,
K. 2009. Type 2C protein phosphatases directly regulate abscisic acid-
activated protein kinases in Arabidopsis. Proceedings of the National
Academy of Sciences of the United States of America 106, 17588–17593.
Vahisalu, T., Kollist, H., Wang, Y.F., Nishimura, N., Chan, W.Y., Valerio, G.,
Lamminmäki, A., Brosché, M., Moldau, H., Desikan, R., Schroeder, J.I.,
and Kangasjarvi, J. 2008. SLAC1 is required for plant guard cell S-type
anion channel function in stomatal signaling. Nature 452, 487–491.
Vahisalu, T., Puzõrjova, I., Brosché, M., Valk, E., Lepiku, M., Moldau, H.,
Pechter, P., Wang, Y.S., Lindgren, O., Salojärvi, J., Loog, M., Kangasjarvi,
J., and Kollist, H. 2010. Ozone-triggered rapid stomatal response
involves the production of reactive oxygen species, and is controlled by
SLAC1 and OST1. Plant Journal 62, 442–453.
Vainio, H., and Bianchini, F. 2003. Fruit and Vegetables. IARC Handbooks of
Cancer Prevention. World Health Organization, Geneva.
Van Doorn, W.G. 1997. Water relations of cut flowers. Horticultural Reviews
18, 1–85.
Van Doorn, W.G. 2012. Water relations of cut flowers: an update. Horticultural
Reviews 40, 55–106.
van Meeteren, U. 1978. Water relations and keeping-quality of cut gerbera
flowers 2. Water balance of ageing flowers. Scientia Horticulturae 9,
van Meeteren, U. 1989. Water relations and early leaf wilting of cut chrysan-
themums. Acta Horticulturae 261, 129–135.
van Meeteren, U., Rezaei Nejad, A., and Harbinson, J. 2009. Effect of (changes
in) air humidity on transpiration and (adaptation of) stomatal closure of
Tradescantia leaves during water stress. Acta Horticulturae 847, 115–122.
Veierskov, B., and Kirk, H.G. 1986. Senescence in oat leaf segments under
hypobaric conditions. Physiologia Plantarum 66, 283–287.
Velez-Ramirez, A.I., van Ieperen, W., Vreugdenhil, D., and Millenaar, F.F. 2011.
Plants under continuous light. Trends in Plant Science 16, 310–318.
Vitagliano, C., and Hoad, G.V. 1978. Leaf stomatal resistance, ethylene evolu-
tion and ABA levels as influenced by (2-chloroethyl) phosphonic acid.
Scientia Horticulturae 8, 101–106.


Vlad, F., Rubio, S., Rodrigues, A., Sirichandra, C., Belin, C., Robert, N., Leung,
J., Rodriguez, P.L., Laurière, C., and Merlot, S. 2009. Protein phospha-
tases 2C regulate the activation of the Snf1-related kinase OST1 by
abscisic acid in Arabidopsis. Plant Cell 21, 3170–3184.
Waller, J.M., Lenné, J.M., and Waller, S.J. 2001. Plant Pathologist’s Pocketbook.
CABI Publishing, Wallingford, UK.
Wang, P., and Song, C.P. 2008. Guard-cell signaling for hydrogen peroxide
and abscisic acid. New Phytologist 178, 703–718.
Wang, Y., Fu, X.Z., Liu, J.H., and Hong, N. 2011a. Differential structure and
physiological response to canker challenge between ‘Meiwa’ kum-
quat and ‘Newhall’ navel orange with contrasting resistance. Scientia
Horticulturae 128, 115–123.
Wang, Y., Li, L., Ye, T., Zhao, S., Liu, Z., Feng, Y.Q., and Wu, Y. 2011b. Cytokinin
antagonizes ABA suppression to seed germination of Arabidopsis by
down regulating ABI5 expression. Plant Journal 68, 249–261.
Wardlaw, C. 1936. Studies in tropical fruits. II. Observations on internal gas
concentrations in fruit. Annals of Botany 50, 655–676.
Waterland, N.L., Campbell, C.A., Finer, J.J., and Jones, M.L. 2010. Abscisic
acid application enhances drought stress tolerance in bedding plants.
HortScience 45, 409–413.
Weaver, L.M., and Amasino, R.M. 2001. Senescence is induced in individually
darkened Arabidopsis leaves, but inhibited in whole darkened plants.
Plant Physiology 127, 876–886.
Wigger, J., Phillips, J., Peisker, M., Hartung, W., Zur Nieden, U., Artsaenko,
O., Fiedler, U., and Conrad, U. 2002. Prevention of stomatal closure by
immunomodulation of endogenous abscisic acid and its reversion by
abscisic acid treatment: Physiological behaviour and morphological fea-
tures of tobacco stomata. Planta 215, 413–423.
Wilkinson, S., and Davies, W.J. 2002. ABA-based chemical signaling: The co-
ordination of responses to stress in plants. Plant, Cell and Environment
25, 195–210.
Wilkinson, S., Clephan, A.L., and Davies, W.J. 2001. Rapid low temperature-
induced stomatal closure occurs in cold-tolerant Commelina communis
leaves but not in cold-sensitive tobacco leaves, via a mechanism that
involves apoplastic calcium but not abscisic acid. Plant Physiology 126,
Williams, M., Brown, M.A., Vesk, M., and Brady, C. 1994. Effect of posthar-
vest heat treatments on fruit quality, surface structure, and fungal dis-
ease in Valencia oranges. Animal Production Science 34, 1183–1190.
Williamson, V.G., and Milburn, J.A. 1995. Cavitation events in cut stems kept
in water: Implications for cut flower senescence. Scientia Horticulturae
64, 219–232.
Willmer, C., and Johnston, W. 1976. Carbon dioxide assimilation in some aer-
ial plant organs and tissues. Planta 130, 33–37.

Stom ata a n d P ostharvest P hy s i o l o g y

Wingler, A., Von Schaewen, A., Leegood, R.C., Lea, P.J., and Quick, W.P. 1998.
Regulation of leaf senescence by cytokinin, sugars, and light effects on
NADH-dependent hydroxypyruvate reductase. Plant Physiology 116,
Wise, R., Olson, A., Schrader, S., and Sharkey, T. 2004. Electron transport is
the functional limitation of photosynthesis in field grown pima cotton
plants at high temperature. Plant, Cell and Environment 27, 717–724.
Xu, Z., and Zhou, G. 2008. Responses of leaf stomatal density to water sta-
tus and its relationship with photosynthesis in a grass. Journal of
Experimental Botany 59, 3317–3325.
Xu, Z.J., Nakajima, M., Suzuki, Y., and Yamaguchi, I. 2002. Cloning and char-
acterization of the abscisic acid-specific glucosyltransferase gene from
adzuki bean seedlings. Plant Physiology 129, 1285–1295.
Xue, H.W., Chen, X., and Mei, Y. 2009. Function and regulation of phospho-
lipid signaling in plants. Biochemical Journal 421, 145–156.
Yakushiji, H., Morinaga, K., and Nonami, H. 1998. Sugar accumulation and
partitioning in Satsuma mandarin tree tissues and fruit in response to
drought stress. Journal of the American Society for Horticultural Science
123, 719–726.
Yan, F., Sun, Y., Song, F., and Liu, F. 2012. Differential responses of stomatal
morphology to partial root-zone drying and deficit irrigation in potato
leaves under varied nitrogen rates. Scientia Horticulturae 145, 76–83.
Yoshida, R., Hobo, T., Ichimura, K., Mizoguchi, T., Takahashi, F., Aronso, J.,
Ecker, J.R., and Shinozaki, K. 2002. ABA-activated SnRK2 protein kinase
is required for dehydration stress signaling in Arabidopsis. Plant and
Cell Physiology 43, 1473–1483.
Yoshida, R., Umezawa, T., Mizoguchi, T., Takahashi, S., Takahashi, F., and
Shinozaki, K. 2006. The regulatory domain of SRK2E/OST1/SnRK2.6
interacts with ABI1 and integrates abscisic acid (ABA) and osmotic
stress signals controlling stomatal closure in Arabidopsis. Journal of
Biological Chemistry 281, 5310–5318.
Zeevaart, J.A.D. 1974. Levels of ({+/−}) abscisic acid and xanthoxin in spinach
under different environmental conditions. Plant Physiology 53, 644–648.
Zeiger, E. 2000. Sensory transduction of blue light in guard cells. Trends in
Plant Science 5, 183–185.
Zeng, W., Melotto, M., and He, S.Y. 2010. Plant stomata: a checkpoint of host
immunity and pathogen virulence. Current Opinion in Biotechnology 21,
Zhang, K., and Gan, S.S. 2012. An abscisic acid-AtNAP transcription factor-
SAG113 protein phosphatase 2C regulatory chain for controlling dehy-
dration in senescing Arabidopsis leaves. Plant Physiology 158, 961–969.
Zhang, K., Xia, X., Zhang, Y., and Gan, S.S. 2012. An ABA-regulated and Golgi-
localized protein phosphatase controls water loss during leaf senes-
cence in Arabidopsis. Plant Journal 69, 667–678.


Zhang, X., Takemiya, A., Kinoshita, T., and Shimazaki, K.I. 2007. Nitric oxide
inhibits blue light-specific stomatal opening via abscisic acid signaling
pathways in Vicia guard cells. Plant and Cell Physiology 48, 715–723.
Zhang, X., Wang, H., Takemiya, A., Song, C.P., Kinoshita, T., and Shimazaki,
K. 2004. Inhibition of blue light-dependent H+ pumping by abscisic acid
through hydrogen peroxide-induced dephosphorylation of the plasma
membrane H+ -ATPase in guard cell protoplasts. Plant Physiology 136,
Zhao, X., Qiao, X.R., Yuan, J., Ma, X.F., and Zhang, X. 2012. Nitric oxide inhib-
its blue light-induced stomatal opening by regulating the K+ influx in
guard cells. Plant Science 184, 29–35.
Zhou, R., Cutler, A.J., Ambrose, S.J., Galka, M.M., Nelson, K.M., Squires,
T.M., Loewen, M.K., Jadhav, A.S., Ross, A.R., and Taylor, D.C. 2004. A
new abscisic acid catabolic pathway. Plant Physiology 134, 361–369.
Zhu, M., Dai, S., and Chen, S. 2012. The stomata frontline of plant interac-
tion with the environment—Perspectives from hormone regulation.
Frontiers in Biology 7, 96–112.
Zhu, S.Y., Yu, X.C., Wang, X.J., Zhao, R., Li, Y., Fan, R.C., Shang, Y., Du, S.Y.,
Wang, X.F., Wu, F.Q., Xu, Y.H., Zhang, X.Y., and Zhang, D.P. 2007. Two
calcium-dependent protein kinases, CPK4 and CPK11, regulate abscisic
acid signal transduction in Arabidopsis. Plant Cell 19, 3019–3036.
Zieslin, N. 1989. Postharvest control of vase life and senescence of rose flow-
ers. Acta Horticulturae 261, 257–264.
Ziv, M., Schwartz, A., and Fleminger, D. 1987. Malfunctioning stomata in
vitreous leaves of carnation (Dianthus caryophyllus) plants propagated
in vitro: Implications for hardening. Plant Science 52, 127–134.

Chapter 7

Water Loss from

Harvested Horticultural
Mikal E. Saltveit
University of California, Davis, California

Abstract 218
7.1 Importance of Water Loss 219
7.2 Properties of Water 220
7.3 Psychrometrics: Behavior of Water in Air 221
7.4 Transpiration: Diffusion of Water Vapor 224
7.5 Resistance to Diffusion of Water Vapor 226
7.6 Measurement of Transpiration 226
7.6.1 Weight Loss 227
7.6.2 Direct Measurement 227
7.6.3 Diffusion Porometer 227
7.7 Factors Affecting Water Loss 227
7.7.1 Commodity Factors 227 Surface-to-Volume Ratio 228 Routes of Water Loss 228 Anatomy of the Evaporating Surface 229 Physiological State of the Commodity 230

P OST H AR V EST RI P ENING P H Y SIOLOG Y Cultivar 230 Cultural Conditions 231
7.7.2 Environmental Factors 231 Humidity 231 Diffusion Shells and Air Velocity 231 Temperature 232 Atmospheric Pressure 232
7.8 Methods to Reduce Water Loss 232
7.8.1 Handling Techniques 232
7.8.2 Proper Refrigeration Design 232
7.8.3 Packaging 233
7.8.4 Waxing 234
7.8.5 Film Wraps 234
7.8.6 Curing 234
7.9 Conclusions and Future Perspectives 235
References 235

Water loss from harvested horticultural commodities will continue to be
a problem as long as warm products need to be cooled and kept cold
during storage and marketing. Even a small amount of water loss can
adversely affect product quality, marketability, and storability. Apart
from the loss of fresh weight that accompanies water loss, a few percent
water loss can stimulate physiological changes that hasten senescence.
The physics of heat transfer in a mechanical refrigeration system neces-
sitates that the air in a cold room will have a vapor pressure less than
that of the commodity. This inherent difference in the vapor pressure
between water near the surface of fresh fruits, vegetables, and ornamen-
tals and the ambient cooling air is the prominent force that determines
the rate of water loss. Increasing the relative humidity of the surround-
ing air or the resistance of the surface of the commodity to the diffu-
sion of water vapor will decrease the rate of water loss. This can be done
by applying a wax or edible coating to the surface of whole or fresh-cut
commodities, by enclosing them in a package that raises the ambient
humidity, and by using harvest and postharvest practices that reduce
injuries (e.g., bruises, wounds, abrasions, etc.) that decrease the ability
of the surface to retard water loss. Preharvest factors (e.g., cultivar selec-
tion, growing environment, cultural practices, and maturity at harvest)
can have significant effects on rates of water loss. Familiarity with the


psychrometric chart provides insights into the relationships among vapor

pressures, relative humidity, and product temperature. Rapid cooling of
warm-harvested commodities and maintaining a high relative humidity
in the cold room is essential to minimize the initial and subsequent level
of water loss.

7.1  Importance of Water Loss

Since most fresh fruits, vegetables, and ornamentals are 90%–95% water,
even a few percent loss of water can produce a significant change in their
postharvest behavior (Table 7.l). Water loss at levels much below those
that produce visible symptoms can influence many of the physical and
physiological changes occurring in harvested horticultural crops. These
effects range from physical (e.g., wilting, shriveling) and economic (e.g.,
weight loss) changes, to subtle visual and physiological (e.g., softening,
senescence) changes. While most effects are deleterious (e.g., reduction of
wound healing in roots and tubers), some may be beneficial (e.g., reduction
of chilling injury symptoms in some sensitive fruit).
Water loss can be a major source of economic loss during the post-
harvest period. The quality losses associated with these changes can result

Table 7.1  Effects of Water Loss on the Postharvest Behavior of Harvested

Horticultural Commodities
Water Loss (%) Possible Change in Postharvest Behavior
0.5 Greater activity of pectolytic enzymes (e.g.,
1.0 Increased CO2 and C2H4 production; faster ripening,
yellowing, and abscission. Reduced wound healing
(periderm formation).
2.0 Reduced turgor, increased ABA content; reduced
susceptibility to chilling injury. Accelerated loss of
3.0 Reduced severity of certain physiological disorders such
as senescent breakdown. Loss of membrane integrity.
4.0 Faster loss of vitamins A and C, and flavor. Discoloration
of mechanical injuries.
5.0 Loss of color intensity and gloss. Accentuation of pitting
associated with chilling injury. Wilting and shriveling.
6.0 Loss of textural quality, e.g., softening, limpness,
flaccidity, and loss of crispness and juiciness.


from a reduction in grade, quality, or price of the commodity. In addition,

the loss of water reduces fresh weight, which directly becomes a reduction
in value for products sold by weight. Water loss may also reduce the total
package weight so that the package may be rejected as no longer meet-
ing the grade standard. Repacking product to meet grade standards can be
very expensive.
An important function of water is to maintain the outward form of
perishable products. The cells in a commodity are like tiny balloons that
are osmotically inflated with water under pressure. The cell wall constrains
this turgor pressure, producing rigid, inflexible cells that contribute to the
texture and appearance of tissues. Water loss produces a reduction in tur-
gor pressure and the shrinkage of cells that becomes visually apparent as
wilting of leafy vegetables, flaccidity of other vegetables, and shriveling and
wrinkling of fruits.
Water loss can also be indirectly responsible for deterioration during
transport and marketing. Many perishables are “bulge” packed so that each
piece is tightly constrained within the box. The inevitable loss of water after
harvest reduces the size of the commodity, and if the individual commodi-
ties are not immobilized by packing material, they can be knocked about
within the package during handling and transport. This can lead to vari-
ous types of transit damage (e.g., scuffing of soft fruits, “roller” bruising of
pears, and shattering of grapes).

7.2  Properties of Water

Water is a remarkable chemical whose properties are fundamental to the
processes of life. The electrical potential surrounding a water molecule is
not uniformly distributed. The partial opposite charges on the oxygen and
hydrogen atoms in a water molecule (i.e., the asymmetric charge separa-
tion that produces polarity) makes water an effective solvent for a wide
range of chemicals, including many biologically important molecules.
Noncovalent hydrogen bonds between the hydrogen of one molecule and
the oxygen of an adjacent water molecule contribute to the exceptional
thermodynamic properties of water. Water has a high heat capacity (i.e.,
it takes a lot of energy to change water’s temperature: 1 kcal/kg/°C) and
an extremely high heat of vaporization (i.e., it takes a lot of energy to turn
liquid water into vapor: 540 kcal/kg/°C). These two properties help sta-
bilize the plant’s temperature (e.g., reducing the rate of freezing on cool
nights, and preventing overheating through evaporative cooling of tissue
exposed to direct sunlight). Water is also incompressible, so it can be an
important structural component of cells.
Water loss from perishable crops results primarily from evaporation
of liquid water from the surfaces of cells and diffusion of this water vapor


away from the tissue. The epidermis of all terrestrial plants is covered with
a waxy cuticle that evolved to significantly reduce diffusion of water vapor
from the tissue. However, this coating also reduces the diffusive exchange
of O2 and CO2 with the external environment. Specialized cells in the epi-
dermis of photosynthetically active leaves (i.e., guard cells) surround a
pore (i.e., the stoma) that breaches the epidermal barrier. Guard cells can
actively change their shape and size to alter the size of the stomatal open-
ing to balance the import of CO2 against the loss of water vapor. In other
organs, static ventilation systems such as lenticels allow diffusion of CO2
and O2 across the epidermis while minimizing water loss. The cuticle may
be modified during growth, development, and ripening to alter its water
diffusion characteristics. The rate of water loss, that is, transpiration, is
therefore controlled by active (i.e., stomata) and passive (e.g., lenticels)
routes of water loss. Transpiration in actively growing plants also provides
the driving force for water uptake by the roots and movement through the
xylem to the leaves.

7.3  Psychrometrics: Behavior of Water in Air

The behavior of water in the vapor phase is termed psychrometrics (i.e.,
properties of water vapor in air at different temperatures). When liquid
water is in contact with air, an equilibrium is established between mole-
cules that evaporate from the liquid’s surface and those that remain in the
liquid phase. At equilibrium, the amount of water vapor in the air above liq-
uid water is dictated by the temperature and the solute concentration. The
contribution of this latter factor is typically ignored since the solute con-
centration of water evaporating from the cell’s surface is usually very low.
In the vapor phase, water vapor acts just like any other gas. The total
pressure of the air in contact with liquid water is usually 1 atm (i.e., 101 kPa,
1013 mbar, 760 mmHg) and is the sum of the partial pressures of the com-
ponent gasses (i.e., N2, O2, Ar, CO2, H 2O, etc.).
The relationship between all the variables involved (see list below)
can be displayed in a psychrometric chart (Figure 7.1). Careful study of
the psychrometric chart reveals a great deal about the behavior of water
molecules in air, which is of some importance to postharvest physiologists
and technologists. Terms used in the psychrometric chart are listed and
defined below:

Dry bulb temperature: The horizontal axis of the chart is the

temperature of a dry thermometer in air.
Vapor pressure (VP): The vertical right-hand axis of the chart
shows the partial pressure of water vapor in the air (e.g., in kPa or
some other measure of pressure).


Relative humidity (%)

100 80 60




Vapor pressure (kPa)

tem 20 2.5



5 1.0
Relative 80 0.5
humidity 40
(%) 20
0 0.0
0 5 10 15 20 25 30 35 40 45
Dry bulb temperature (°C)

Figure 7.1  A psychrometric chart showing the interrelationships among dry

and wet bulb temperatures, vapor pressures, and relative humidity.

Wet bulb temperature: The left-to-right upward curving lines

show the temperature of a thermometer whose bulb is covered
with ventilated water-soaked gauze.
Dew point temperature: The horizontal lines extending from the
right axis and intersecting the wet bulb temperature line at the
dew point, the temperature of a surface upon which liquid water
will condense.
Saturation vapor pressure (SVP): The maximum amount of
water vapor (expressed as a partial pressure) that air at a specific
temperature can hold. It is found by extending a horizontal line
from the wet bulb temperature to the right boundary of the chart.
Relative humidity (RH): The ratio of the actual water content
of the air to the maximum possible water content under any par-
ticular temperature. The RH lines are represented by the equally
spaced curved lines that slope upward from left to right. The RH
can be expressed for a particular air sample and temperature as


RH  =  (VP/SVP) × 100, where RH = percent of maximum water

vapor in the air sample at that temperature, SVP = saturation vapor
pressure at 100% RH and that temperature, and VP = vapor pres-
sure in the air sample at that temperature.

Let’s use a few examples to see how we can use the psychrometric chart
(Figure 7.1).
The relative humidity (RH) can be determined by finding the inter-
section of the line extending upward from the dry bulb temperature with
the downward-sloping line from the wet bulb temperature. For example,
air with a dry bulb temperature of 25°C and a wet bulb temperature of 18°C
has a RH of 50% (point A on Figure 7.2). The vapor pressure (VP) of that
50% RH air can be determined by following the horizontal line to its inter-
section with the vertical axis at 1.6 kPa (point B on Figure 7.2). The dew
point temperature of 14°C can be determined by following the horizontal

Relative humidity (%)

100 80 60




Vapor pressure (kPa)


20 2.5



15 A B
C 14

5 1.0
Relative 80 0.5
humidity 40
(%) 20
0 0.0
0 5 10 15 20 25 30 35 40 45
Dry bulb temperature (°C)

Figure 7.2  Example of how the psychrometric chart can be used to find the
vapor pressure and dew point from measurements of the wet and dry bulb
temperatures (refer to text for detailed description).


line left to its intersection with the wet bulb temperature curve (point C on
Figure 7.2).
In another example, the RH of air with a dry bulb temperature of 15°C
and a wet bulb temperature of 12°C will be 70%, with a vapor pressure of
1.2 kPa and a dew point of 10°C.
If a sample of warm moist air is cooled (move left along a horizontal
line), its relative humidity increases until it reaches saturation (at the left-
most line, the wet bulb temperature curve). The cooled air is now at its dew
point. Further removal of heat will result in condensation of water from the
air onto the cool surface. In contrast, the RH of air rapidly decreases with
increasing temperature (move right along the horizontal line from the wet
bulb temperature curve). This means that at warm temperatures, air can
hold much more water vapor than at cool temperatures. The water holding
capacity of air just about doubles with every 11°C rise in temperature.
If a cold commodity is placed in a warm room, the air in contact with
the commodity will become cooler, and depending on the initial RH of the
air, water may condense on the surface of the fruit. In contrast, if a warm
commodity is placed in a cool atmosphere, the air in contact with the fruit
will become warmer (move to the right along the horizontal lines) and its
RH will decrease, thereby facilitating water evaporation from the tissue.

7.4  Transpiration: Diffusion of Water Vapor

For any gas, the rate at which it diffuses between two points is proportional
to the difference in partial pressures between the points and the diffusive
resistance of the intervening material. The rate at which water vapor dif-
fuses from a commodity is therefore proportional to the difference between
the vapor pressure at the surface of the commodity and the vapor pressure
of the surrounding air. The vapor pressure of water molecules in the air
can be determined from the psychrometric chart for any temperature and
RH. Because air inside a commodity is in equilibrium with the almost pure
water in the cell walls, the vapor pressure in the air spaces within tissue is
close to the saturated vapor pressure at that temperature. The vapor pres-
sure difference between the tissue and the outside air can therefore be cal-
culated with the equation
VPD = SVPinternal − VPexternal
where VPD = vapor pressure difference between the tissue and the
­surrounding air, SVPinternal = saturation vapor pressure of air at the tem-
perature of the tissue, and VPexternal = vapor pressure in the external air at
its temperature and RH.
The VPD can be used to determine the rates of water loss for different
combinations of commodity, air temperature, and RH using the following


equation (these calculations are only correct when other conditions, such
as barometric pressure, nature of the commodity’s surface, and air velocity
past the product remain constant).
J = K × VPD
where J = rate of water loss (g/h, % FW/day, etc.), K = ­proportionality
­constant (a function of many features of the commodity that will be
discussed later), and VPD = vapor pressure difference, or deficit (kPa,
mmHg, mbar, etc.).
For example, consider a harvested head of lettuce at 25°C that loses
1% of its fresh weight when held for 1 hour (i.e., J = 1%/h) in a room at 25°C
and 80% RH. What will be its initial and final rates of weight loss during
cooling to 4°C in a store at 80% RH (Figure 7.3)?
From the psychrometric chart we can determined that at 25°C, the
­saturation VP is 3.2 kPa (point A on Figure 7.3). The VP of the 25°C  air

Relative humidity (%)

100 80 60




Vapor pressure (kPa)


20 2.5

VPD = 3.2 – 0.6 = 2.6




5 1.0
0 B
Relative 80 0.5
VPD = 0.8 – 0.6 = 0.2
(%) 20
0 0.0
0 5 10 15 20 25 30 35 40 45
Dry bulb temperature (°C)

Figure 7.3  Example of how the psychrometric chart can be used to calcu-
late the vapor pressure deficit (VPD) experienced by a head of lettuce placed
into a cold room as it cools from 25°C to 4°C (refer to text for detailed


at  80% RH is 80% of the saturation VP or about 2.5 kPa, and the VPD is
the difference of the two values, or about 0.7 kPa (i.e., 3.2 − 2.5 = 0.7).
Substituting in the equation J = K × VPD, we can calculate the proportional-
ity constant K [K = (1%/h)/(0.7 kPa) = 1.4% FW/kPa-h].
The VP of 100% RH air at 4°C is 0.8 kPa, while 80% RH air at the
same temperature has a VP of 0.6 kPa (0.8 × 0.80) (point B of Figure 7.3).
When the 25°C lettuce is initially put in the 80% RH 4°C cold room, the VPD
between the tissue and the air will be 2.6 kPa (i.e., 3.2 − 0.6). The initial rate
of water loss will therefore be 2.6 kPa × 1.4% FW/kPa-h = 3.6% FW/h. This
initial high rate of water loss is one reason why cooling to the correct stor-
age temperature should be done as quickly as possible. Once the product
is cooled to 4°C, the VP of the lettuce decreases to 0.8 kPa and the VPD
between the product and the air decreases to 0.2 kPa (0.8 − 0.6). The new
rate of water loss is 0.2 kPa × 1.4% FW/kPa-h = 0.28% FW/h. This amounts
to a 92% reduction in the rate of water loss.

7.5  Resistance to Diffusion of Water Vapor

The transpiration equation ( J = K × VPD) is a specific example of Fick’s law
of gas diffusion, which in the case of the diffusion of water vapor states that
J = (Area/Resistance) × (SVPinternal − VPexternal) = (A/R) × (VPD)
where J = rate of water vapor loss, A = surface area over which water vapor
is being lost to the external environment, R = coefficient of resistance to
water vapor diffusion, and VPD = vapor pressure difference, or deficit (kPa,
mmHg, mbar, etc.).
Once VPD is minimized by reducing the temperature and increasing
the RH around the commodity, the resistance to diffusion (R) is the most
important constant for determining susceptibility of different products to
the loss of water vapor. Because R is a resistance coefficient, higher val-
ues indicate lower rates of water loss. The composition of the cuticle, and
its resistance to diffusion, may change in response to growing conditions,
during development, and during postharvest handling. Waxing and film
coatings are some postharvest practices that are used to increase the com-
modities’ resistance to water vapor loss.

7.6  Measurement of Transpiration

Measurements of the rates of water loss are as important to the postharvest
physiologist as are measurements of respiration and ethylene production.
Several methods are commonly used; the most important are described in
the following subsections.


7.6.1  Weight Loss

Since almost all weight loss is due to water loss, the simplest method to deter-
mine the rate of water loss is to periodically weigh the commodity and cal-
culate the rate of water loss over time. In a few cases (e.g., long-term storage
of commodities such as onions), the dry weight loss due to respiration may
contribute to total weight loss and can be a source of errors with this method.

7.6.2  Direct Measurement

Measurement of changes in relative humidity (RH) or dew point of air in
a container enclosing a known weight of commodity can be used to deter-
mine the transpiration rate; this is similar to the method used in a static
or flowing system to measure the rate of respiration (see Chapter 5). This
method may be useful for researchers wishing to study water loss in com-
modities under controlled atmospheres or when CO2 and C2H4 production
is also being measured.

7.6.3  Diffusion Porometer

The diffusion porometer, designed to measure diffusive resistance of leaves
(and thereby to estimate opening of the stomata), can be used to measure
the diffusive resistance of those perishable commodities that lose water
fairly readily.

7.7  Factors Affecting Water Loss

As noted above, the equation used to describe water loss from perishables is,
in fact, a simplified version of an important law governing the diffusion of all
gases, Fick’s law of diffusion. This law states that diffusion of gases across a
surface is governed by the area of the surface, the resistance of the surface to
diffusion, and the concentrations of gases on either side of the surface. The
last term is governed by the temperature and RH in the storage environment,
while the other two terms are dependent on the properties of the commodity.

7.7.1  Commodity Factors

Commodities differ in size, shape, composition, physiology, and the compo-
sition and topology of the epidermis and cuticle. Many of these properties
can be significantly changed due to the conditions under which they were
grown and under which they are handled after harvest. The same cultivar


can have markedly different rates of water loss depending on where and
when it was grown and how it was handled during and after harvest.  Surface-to-Volume Ratio

The rate at which water is lost from a commodity is a function of its sur-
face area. If one calculates water loss as a percentage of the initial product
weight for commodities whose surfaces have the same resistance to water
vapor diffusion, the rate of water loss will be a function of the surface
area-to-volume ratio of each commodity. Simple formulae for calculating
surface area and volume (Table 7.2) of commodities reveal major differ-
ences in the surface-to-volume ratio between large and small commodi-
ties (Table 7.3). If the rate of water loss per unit surface area is identical
for large and small units of a commodity, then the large item will still
lose less water per unit weight than the smaller item because it has less
surface area to volume than the smaller unit. For any commodity, smaller-
sized items will lose water at a faster rate than larger items.  Routes of Water Loss

There are two routes by which perishable commodities lose water.
Epidermal cells on the surface of the commodity can lose water directly
through their surface. The outer surfaces of all plant parts are covered with
water-repellant coverings: cuticle in aerial organs and suberin in roots and

Table 7.2  Equations for the Calculation of the Area and Volume for a Sphere
(Tomato), a Cylinder with Spherical Ends (Cucumber), and A right Circular
Cone with One Hemispherical End (Carrot).
Sphere Cylinder Cone
Area 4πr2 2πLr + 4πr 2 πr'r2 + L2 + 2πr2
Volume 4/3πr 3 πLr2 + 4/3πr3 1/3πLr2 + 2/3πr3

Table 7.3  Area and Volume of Some Commodities

Length Radius Area Volume
Commodity (cm) (cm) (cm2) (cm3 or mL) Area/Volume
Tomato (small) 4 201 268 0.75
Tomato (large) 8 804 2,144 0.38
Cucumber (small)  8 2 151 134 1.12
Cucumber (large) 16 4 603 1,072 0.56
Carrot (small)  8 2  89 50 1.78
Carrot (large) 16 4 358 402 0.89


tubers. Water losses through the epidermal cells occur by a relatively com-
plex process whereby water molecules must dissolve into the epidermal cov-
ering, diffuse through it, and then evaporate into the outside atmosphere.
The rate at which evaporation occurs is often limited by the solubility of
water in the outer surface coating.
The surface coating is very effective in retarding water loss. For exam-
ple, evaporation from the cells inside the leaf is 10-fold higher than from
cells on the leaf surface. In very heavily cutinized or suberized organs, very
little water escapes by direct evaporation from the surface cells. Instead,
most water is lost in the vapor phase through apertures in the surface of
the organ, such as stomata, lenticels, and wounds. The rate of evaporation
is affected not only by the route of evaporation, but also by the architecture
of the evaporating surface.  Anatomy of the Evaporating Surface

Important routes of evaporation and anatomical features affecting its
rate are

Stomata: Specialized organelles common to all green tissues of

higher plants; stomata are designed to facilitate inward diffusion
of CO2 during photosynthesis, but to minimize water loss at times
of low light or during water stress. Typically, the lower surfaces of
leaves are much more richly supplied with stomata than the upper
surface. Stomata will normally close when a tissue or organ is
under water stress, but can remain open during sudden reduction
in temperature (such as occurs during forced-air cooling), thus
aggravating water loss during storage.
Lenticels: In large organs, there is a danger that insufficient O2
will diffuse into the tissues to prevent anaerobiosis because of the
diffusive resistance of the cuticle. Lenticels are growths of unsu-
berized subepidermal cells that erupt through the epidermis and
facilitate the passive ventilation of bulky tissues. Changes in their
structure and thickness in response to environmental changes
during development and after harvest can modify their diffusive
Surface imperfections: In many commodities, especially after
harvest, water escapes through various imperfections in the epi-
dermis. Some of these imperfections may be the result of abnor-
mal growth; others, such as stem scars, open calyx, and blossom
scars, are normal anatomical features. Wounds of any sort also
result in increased water loss. Bruising, abrasion, and most
importantly, actual rupture of the epidermis result in greatly
accelerated water loss.


Cuticular waxes: The chemical composition and a­ rchitecture

of the cuticular surface can dramatically affect water loss.
Thickness of the cuticle obviously affects the rate of cuticular
diffusion of water. Waxes deposited on the surface of maturing
fruits and vegetables frequently have a complex architecture
composed of randomly oriented plates and fibers. These obstruc-
tions to airflow produce diffusion shells above the surface that
retard water loss.
Trichomes: Water loss in many plant parts is reduced by the pres-
ence of surface appendages called trichomes. These may be single
celled, multicelled, tufted, star shaped, or branched. Although
trichomes increase the surface area of an organ, they also trap
saturated water vapor around it and reduce air velocities past the
evaporating surface. Breakage of hairs or trichomes in posthar-
vest handling can aggravate water loss from commodities such as
peaches, kiwifruit, and zucchini.
Architecture: The effect of the surface-to-volume ratio on the
rate of water loss has been discussed already. In many plant
parts, the way in which the organ is constructed may also have
a dramatic effect on the rate of water loss. A striking example
is the difference between leaf lettuce and head cabbage. Both
are leafy vegetables, but the condensed cabbage offers little of
its leaf surface to evaporation. In compact leafy vegetables such
as cabbage or onion, the rate of water loss may be limited by the
ability of the vascular system to provide water to the outer evapo-
rating leaves. The desiccated scale leaves of onion or wrapper
leaves of cabbage can further reduce the rate of water loss from
the commodity.  Physiological State of the Commodity

It is obvious that young vegetative tissues, such as watercress, with plen-
tiful stomata and thin cuticles, lose a great deal more water than heavily
suberized storage organs such as potato. More subtle changes in physiolog-
ical state, such as ripening, can also strongly affect the rate of water loss.
The effect of ripening on the composition and structure of the surface wax
of apples has been alluded to before. A similar effect is probably responsible
for the difference in the water loss from dark green and yellow lemons. Cultivar
The many structural and chemical differences between different cultivars
can substantially affect weight loss.

WATER LOSS  Cultural Conditions

Seasonal differences in environmental conditions during the growing
period may also affect the rate of weight loss from harvested commodities.
The porosity and diffusivity of the epidermis and cuticle can vary signifi-
cantly between the same commodity grown in hot versus cool and humid
versus dry environments. There are even differences in the rate of water
loss from fruit grown in the shaded inside of the tree and those grown on
the sunny periphery of the tree.

7.7.2  Environmental Factors

The postharvest environment can have a substantial effect on the rate of
water loss from harvested commodities. Environmental factors are particu-
larly important during long-term storage. Humidity
The importance of the water vapor content of the air on water loss has
been emphasized already. Lower-humidity environments also frequently
adversely affect the quality of stored commodities. Because of the natural
cycling of temperatures in cold storage rooms, a compromise must be made
between the danger of condensation of water on cold fruit during the warm
part of the cycle in rooms with very high RH, and unacceptable levels of
water loss if the RH is maintained at levels safe for the entire cycle.  Diffusion Shells and Air Velocity

When water vapor diffuses beyond the epidermis, it has not yet entirely
escaped from the influence of the commodity. Thin surface layers of satu-
rated air, termed diffusion shells, which may be pronounced in commodi-
ties with hairs or elaborately sculptured wax deposits on their surface,
provide a significant further barrier to diffusion of water vapor. For this
reason, the rate at which the ambient air is moving past the commodity can
strongly affect the rate of water loss.
As the air velocity increases, the rate of water loss increases in a
hyperbolic fashion, eventually reaching a plateau when, presumably, the
diffusion shells have been completely swept away. In the design of cooling
and storage facilities, and in evaluating postharvest handling systems, it is
important to choose an appropriate compromise between air velocities that
will maintain adequate temperature control and those that will exacerbate
the loss of water from the product.

The close interdependence of temperature, VP, and RH is implicit in the
psychrometric chart, and the importance of temperature in the VPD-driven
loss of water from perishable commodities cannot be overemphasized.
Some studies have shown a substantial effect of temperature on water loss,
even allowing for the changes in VPD. Enhanced respiration and changes
in the composition of the cuticle at warmer temperatures may account for
these observations.  Atmospheric Pressure

Reducing the total atmospheric pressure (e.g., at high altitudes or under
hypobaric conditions during storage or transport) reduces the partial pres-
sure of water vapor and other gases in the air. This, in turn, accelerates
water loss (outward diffusion of water vapor) from fresh produce. This is
the basis of vacuum cooling, where a lowered pressure facilitates evapora-
tive cooling of the commodity.

7.8  Methods to Reduce Water Loss

A wide range of tools are available to the postharvest technologist for reduc-
ing the rate of water loss from perishable commodities. These techniques
are designed to reduce water loss either by reducing VPD or by increasing
the resistance of the surface-to-water diffusion.

7.8.1  Handling Techniques

The primary factors in reducing water loss are the care and rapidity with
which the commodity is removed from the plant, packed, and brought to
the optimum storage temperature. Rough handling will increase surface
damage and water loss through the ruptured epidermis. Tardiness in han-
dling, packing, and cooling will increase the time over which the product is
exposed to high VPDs and result in high water loss.

7.8.2  Proper Refrigeration Design

It is desirable to maintain high relative humidity in the cold room, especially
for the long-term storage of perishables. In general, the highest humidities
are obtained by jacketing the storeroom to reduce heat infiltration and loss
of humidity. There should be more than enough refrigeration capacity to


remove heat from all anticipated sources. Evaporators should be as large as

possible to permit the removal of the required amount of heat without having
to be run at a large temperature differential. The transfer of heat from the
storage room air to the cold evaporator coils is governed by the temperature
differential between them and the volume of air passing over the coils. If the
evaporator is too small, a large temperature differential will be needed to
transfer enough energy to maintain the storage temperature. A very low coil
temperature will be needed to produce the needed large temperature dif-
ferential. Water will condense on the cold coils, thereby lowering the water
content of the cold air exiting the evaporator. As the air warms to the storage
room temperature, its relative humidity will fall to undesirable levels.
In recent years, much has been claimed about the “humifresh” sys-
tems, in which the storage room air is circulated through baths or curtains
of water at or near 0°C. These systems allow very high humidity and very
uniform temperatures, but can be limited in their capacity and ability to
maintain temperatures at or near freezing.
Humidifiers raise the humidity by injecting mist or fog into the stor-
age room, but unless the refrigeration plant is designed to accommodate
the extra load of condensate, the evaporator coils may become covered
with ice, resulting in an actual reduction in humidity due to lowered coil
temperatures. Modern equipment can be fitted with hot gas bypass valves,
which allow the surplus capacity of the refrigeration system to bypass the
condenser when the commodities have attained store temperature and the
refrigeration load is lighter. This system can also divert warm gas to peri-
odically defrost the evaporator coils.
The importance of air velocity to weight loss was noted above. Storage
rooms should be designed to maintain proper product temperatures without
excessive rates of air movement. High air velocities are often used initially
to rapidly cool produce (e.g., forced-air precooling). Thereafter, convective
systems are usually adequate to maintain temperatures and will prevent
excessive weight loss.

7.8.3 Packaging
Placing commodities in packages of any sort obviously changes their geom-
etry with respect to weight loss. The larger the package, the smaller is its
surface area in relation to its volume. This is because the surface area
increases as a function of the dimensions of the package squared, while
the volume increases as a function of the dimensions cubed. In effect, the
package reduces the VPD within the package because the air inside the
package will be at high RH. To further reduce water loss, some products
are packed in polyethylene or other plastic liners, usually ventilated to avoid
anaerobic or high CO2 conditions in the package. Airflow through holes in


the package to maintain the desired product temperature and gas concen-
trations will nullify this benefit.
Certain containers, such as wooden or plain fiberboard boxes, can
absorb a substantial amount of water, thereby decreasing the initial VP in
the package. It is not feasible to equilibrate the containers with the storage
atmosphere, and waxed fiberboard cartons may be too expensive for many
commodities. Under these circumstances, waxed paper or polyethylene lin-
ers can substantially reduce weight loss for little extra cost.

7.8.4 Waxing
An alternative route to those noted above for reducing water loss is to raise
the resistance of the fruit surface to diffusion of water vapor by the applica-
tion of wax. Waxing is widely used as a treatment for fruits, frequently for
cosmetic reasons rather than to reduce weight loss. The waxes employed
are proprietary formulations and probably comprise largely waxes derived
from petrochemicals. Remember that although waxing can slightly reduce
the rate of water loss, temperature (even allowing for differences in the
VPD) can have a far greater effect than waxing. Waxing can also modify
the  internal atmosphere of perishable commodities. There is a danger
that the use of surface coatings can produce toxic internal levels of CO2,
­particularly when the commodity is returned to ambient temperatures after
transportation or storage.

7.8.5  Film Wraps

In recent years, there has been much interest in the use of individual film
wraps for packaging and marketing some fruits and vegetables. This tech-
nology has become a commercial proposition with grapefruit and lemons.
Shrink-wrapping individual grapefruit in high-density polyethylene film
reduced the rate of water loss from the fruits. This technique also reduces
the occurrence of decay and several disorders common in harvested citrus.
The relative contributions of reduction in water loss, isolation of individual
fruits, and modification of the fruit’s internal atmosphere to this beneficial
response have not yet been determined.

7.8.6 Curing
In some commodities, the formation of new or heavier deposits of suber-
ized cells on the surface will substantially increase their resistance to water
loss. Onions, potatoes, and sweet potatoes are frequently cured to improve


their postharvest life and reduce water loss. High temperatures and RH
often assist in the suberization process that occurs during curing.

7.9  Conclusions and Future Perspectives

Reducing water loss is an important consideration in deciding which post-
harvest practices are used. Understanding how the physical laws of gas
diffusion govern water loss and how the concepts visualized in the psychro-
metric chart can be used to calculate the rate of water loss is essential to
formulating and implementing postharvest practices that will minimize
the detrimental effects of water loss. Future practices and technologies to
minimize water loss must be devised to act within the constraints imposed
by these physical laws and also within government regulations. Consumer
preferences will continue to identify those factors that must be either mini-
mized or maximized to optimize product quality. Economic considerations
are also important since no practice will be widely adopted unless it offers a
significant economic advantage over existing systems. The current desire
to consume more locally grown food may minimize the importance of
reducing the rate of water loss since the time from harvest to consumption
would be reduced. In contrast, the desire for more exotic horticultural com-
modities, the desire that seasonal crops be available year-round, and the
basic economics of large-scale production may all contribute to increase
transit times (and levels of water loss) as the distance between areas of pro-
duction and consumption increases. Future research should study package
designs that minimize water loss and cuticular and epidermal properties
that maximize the resistance to water vapor diffusion.

Kader, A.A. 2002. Postharvest Technology of Horticultural Crops. 3rd ed.
Division of Agriculture and Natural Resources, University of California,
Kader, A.A., and Saltveit, M.E. 2003. Respiration and gas exchange. In Bartz,
J.A., and Brecht, J.K. (eds.), Postharvest Physiology and Pathology of
Vegetables. Marcel Dekker, New York, pp. 7–30.
Kays, S.J., and Paull, R.E. (eds.). 2004. Postharvest Biology. Exon Press,
Athens, GA.
Wills, R.B.H., McGlasson, B., Graham, D., and Joyce, D. (eds.). 2007.
Postharvest: An Introduction to the Physiology and Handling of Fruit,
Vegetables and Ornamentals. CAB International, Wallingford, UK.
Woods, J.L. 1990. Moisture loss from fruits and vegetables. Postharvest News
and Information 1, 195–199.

Chapter 8

Lysophospholipids and
Postharvest Quality
of Fruits, Vegetables,
and Cut Flowers
Domingos P.F. Almeida
Universidade de Lisboa, Lisbon, Portugal

Abstract 238
8.1 Introduction 238
8.2 Lipids as Signal Molecules in Plant Senescence and
Stress Response 239
8.2.1 Chemistry 239
8.2.2 Metabolism 240
8.3 Modulation of Postharvest Quality by Lysophospholipids 241
8.3.1 Effects on Color 242
8.3.2 Effects on Texture 242
8.3.3 Other Effects on Postharvest Quality-Related
Features 246
8.4 LPE Treatment Condition 247
8.5 Improvement of Postharvest Quality by
Lysophospholipids: Potential and Limitations 248
8.6 Conclusions and Future Perspectives 249
References 249


Regulatory lipid molecules are involved in plant responses to ­environmental
stresses and developmental cues. Phosphatidic acid is one of the best-
known signaling lipids in plants, but other lipidic molecules are involved in
signal transduction pathways. Therefore, the potential exists for the tech-
nological or biotechnological use of signaling lipids as tools to modulate the
features of fresh produce with postharvest relevance. One attempt has been
made with the exogenous application of lysophosphatidylethanolamine
(LPE) to crops and produce. Modulation of postharvest quality by lyso-
phospholipids has been documented in dozens of horticultural commodi-
ties. Reported effects include delayed senescence and improved color and
texture, among other postharvest quality-related features. Despite the evi-
dence for the regulatory effects of lysophospholipids, two decades of trials
of LPE applications to horticultural produce have not yet been conclusive
regarding the development of lipids as plant growth regulators to improve
postharvest quality. Different approaches can be envisioned to harness the
potential for quality modulation by regulatory lipids: genetic engineering
to alter endogenous levels of regulatory lipids, pharmacological treatments
to target the biosynthesis of the signal transduction pathways of regulatory
lipids, and exogenous application of formulations containing signaling or
regulatory lipids. The improvement of our fundamental understanding of
lipid-mediated signaling and metabolic regulation is likely to result in novel
technological applications aimed at modulated produce quality.

8.1 Introduction
The importance of membranes on the postharvest quality of perishable
plant organs has long been recognized for their structural role, cellu-
lar compartmentation, and site of intense metabolic activity (Marangoni
et al., 1996). However, in addition to these functions, lipids have emerged
as an important class of regulatory and signaling molecules, and their role
in responses to environmental stresses and developmental cues is now
evident in plants and mammalians. Many regulatory lipids are related to
membrane phospholipids, as anabolic precursors or degradation products.
Phosphatidic acid is one of the best-known signaling lipids in plants (Wang
et al., 2006), but other lipidic molecules are involved in signal transduction
pathways. Therefore, the